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SMITHSONIAN INSTITUTION 
UNITED STATES NATIONAL MUSEUM 


PROCEEDINGS 


OF THE 


UNITED STATES NATIONAL MUSEUM 


VOLUME 103 
NUMBERS 3311-3337 


PRAE- INCRE 
= BIE Vice: 4 
PEERY 


UNITED STATES 
GOVERNMENT PRINTING OFFICE 
WASHINGTON : 1956 


ADVERTISEMENT 


The scientific publications of the National Museum include two 
series, known, respectively, as Proceedings and Bulletin. 

The Proceedings, begun in 1878, are intended primarily as a medium 
for the publication of original papers, based on the collections of the 
National Museum, that set forth newly acquired facts in biology, 
anthropology, and geology, with descriptions of new forms and 
revisions of limited groups. Copies of each paper, in pamphlet 
form, are distributed as published to libraries and scientific organiza- 
tions and to specialists and others interested in the different subjects. 

The dates at which these separate papers are published are recorded 
in the tables of contents of each of the volumes. 

The present volume is the hundred and third of this series. 

The Bulletin, the first of which was issued in 1875, consists of a 
series of separate publications comprising monographs of large 
zoological groups and other general systematic treatises (occasionally 
in several volumes), faunal works, reports of expeditions, catalogs of 
type specimens, special collections, and other material of similar 
nature. The majority of the volumes are octavo in size, but a quarto 
size has been adopted in a few instances in which large plates were 
regarded as indispensable. In the Bulletin series appear volumes 
under the heading Contributions from the United States National 
Herbarium, in octavo form, published by the National Museum since 
1902, which contain papers relating to the botanical collections of the 
Museum. 

REMINGTON KELLOGG, 
Director, United States National Museum. 


CONTENTS 


Pages 
Assort, R. Tucker. Review of the Atlantic periwinkles 


Nodilittorina, Echininus, and Tectarius. Figures 55-57. 

Published March 23, 1954... . . 449-464 
Arnett, Ross H., Jr. Beetles of the Soaeitetd wenn 

Vasaces Glasiaanppitee Figure 13. Published April 30, 


Sees CAD OGAROMONI\, BUS a SAI LID. HBR EG POLO, & 19. engyeod 
New species: Vasaces linearis, V. knulli, V. maculatus, V. elon- 
gatus. 


A review of the beetle family Cephaloidae. 
Figure 20 and plate 5. Published May 15, 1953 . . . . 155-161 
Supplement and corrections to J. A. Hyslop’s 

Genotypes of the elaterid beetles of the world. Published 

April 14,1955 . . Po AR CUREe A165 
BANNER, oe H. W pelermeat a W. M. Tattersall’s 

Review of the Mysidacea of the United States National 

Museum. Published July 8, 1954... .. . 575-583 
Bercer, ANDREW J. On the anatomy and Foldatatiins 

of glossy cuckoos of the genera Chrysococcyr, Lampro- 

morpha, and Chalcites. gsi 69-71. Published January 

J Gs LOLS A ee ee ee OOO. 
BuakeE, Doris Hatiunie The chrysomelid beetles of the 

genus Strabala chevrolat. said 17. Published June 5, 

Lis parisianat ; : : . 121-134 


New species: Strabala nai sg. een s. shui, S. 
trinitatis. 

New subspecies: Strabala ambulans jamaicensis, S. a. puerto- 
ricensis, S. rufa floridana, S. acuminata teapensis, S. a. costa- 
ricensis. 


Carriker, M. A., Jr. Studies in Neotropical Mallophaga, 
XI: Bird lice of the suborder Amblycera, genus Dennyus 
Neumann. Figures 63, 64. Published May 21, 1954 . 533-549 
New species: Dennyus brevicapitis, D. intonsus, D. rotundocapitis, 
D. brunneitorques, D. similis, D. spininotus, D. limbus. 
Cartwricut, O. L. Scarabaeid beetles of the genus 
Bradycinetulus and closely related genera in the United 
States. Figures 14-16 and plates 3, 4. Published 
eos fogs, . 5) tere Te eee eke 
New genera: Bolborhombus, Bulbvbescansk 
New species: Bradycinetulus rex, Bolbocerastes regalis, B. impert- 


alis, Bolborhombus parvulus. 
New subspecies: Bolbocerastes imperialis kansanus. 


Tit 


IV PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


CarvatHo, José OC. M. Neotropical Miridae, LXIV: vas 
New bugs of the subfamily Cylapinae (Hemiptera). 
Figures 72-76 and plate 15. Published March 2, 1955 . 621-632 

New genera: Brachyfulvius, Peritropoides. 

New species: Brachyfulvius chapini, Peritroipoides annulatus, 
P. quadrinotatus, Vannius oculatus, Fulvius albonotatus, F. 
castaneous, F. ornatifrons. 

Gurney, Asuitry B. Notes on the biology and immature 
stages of a cricket parasite of the genus Rhopalosoma. 

Figures 8, 9 and plate 1. Published March 10,1953. . 19-34 

Distribution, general bionomics, and recogni- 

tion characters of two cockroaches recently established 

in the United States. Figure 10 and plate 2. Pub- 


lished March 10, 1953. vier: Se Ae 39-56 
Hersuxovirz, Pure. Mammals of northern Colombia, 

preliminary report No. 7: Tapirs (genus Tapirus), with 

a systematic review of American species. Figures 58- 

62. Published May 18, 1954 465-496 


Hyman, Lissre H. North American triclad Turbellaria, 
XIII: Three new cave planarians. Figures 65-68. 
Published June 14,.1954.. 0. sane am saeht “lor yee ees 


New species: Phagocata cavernicola, Sphalloplana georgiana, 
Speophila hoffmastert. 


Kenx, Roman. The fresh-water triclads (Turbellaria) of 
Alaska. Figures 21-25 and plates 6-8. Published 
Fume 5, 1958. cies. osc Wine «Saher s <0 te gels ee 


New species: Phayocata nivea, Polycelis borealis, Dendrocoelopsis 
piriformis, D. alaskensis. 


Lacuner, Ernest A. A revision of the goatfish genus Upe- 
neus with descriptions of two new species. Plates 13, 14. 
Published, May 18, 1954....< 00% as aeyitem+ sah <0: Tet een 


New species: Upeneus asymmetricus, U. oligospilus. 


La Rivers, Ira. Two new naucorid bugs of the genus 
Ambrysus. Figure 1. Published February 12, 1953. . 1-7 


New species: Ambrysus thermarum, A. bispinus. 


LAwReENcE, R. F. Two new scale-mite parasites of lizards. 
Figures 2-7. Published March 10,1953... ... . 9-18 


New species: Geckobia keegani, G. philippinensis. 


McDermort, Frank A. Photuris bethaniensis, a new lam- 
pyrid firefly. Published February 26,1953... ... 35-37 


New species: Photuris bethaniensis. 


CONTENTS 


Morrison, J. P. E. The relationships of Old and New 
World melanians. Plate 11. Published April 20, 1954 . 
Perriponge, Martan H. Marine polychaete worms from 
Point Barrow, Alaska, with additional records from the 
North Atlantic and North Pacific. Figures 26-39. Pub- 
lished May 21, 1954. 2 
ScHULTZ, eeeien P. Review 3 Te finde Baiae anemone 
fishes, genus Amphiprion, with descriptions of two new 
species. Plates 9,10. Published July 3, 1953 . 
New species: Amphiprion tricinctus Schultz and Welander, Am- 
phiprion mauritensis Schultz. 
AND Marsa, N. B. A review of the labrid fish 
genus Wetmorella with descriptions of new forms from the 
tropical Indo-Pacific. Figures 52-54 and plate 12. Pub- 
lished April 21, 1954. . E 


New species: Wetmorella ocellata, W. albofasciata, W. triocellata. 
New subspecies: Wetmorella philippina bifasciata. 


Srurtevant, A. H. Nearctic flies of the family Periscelidae 
(Diptera) and certain Anthomyzidae referred to the family. 
Published June 16, 1954 . Fat bad 


New species: Periscelis occidentalis, Cyamops imitata. 


Waitt, Turopore E. Preliminary analysis of the fossil 
vertebrates of the Canyon Ferry Reservoir area. Figures 
40-51. Published May 28, 1954 . 

New genus: Kentrogomphios. 
New species: Kentrogomphios strophensis, Paradjidaumo_ spo- 
kanensis, Eumys cricetodontoides, E. latidens, E. spokanensis. 

Wirtn, Wiis W. Biting midges of the heleid genus Stilo- 
bezzia in North America. Figures 11, 12. Published 
May 15, 1953. . : 

New species: Stilobezzia (Neostilobezzia) stonet, S. (S.) beckae, S. 
(S). punctipes, S. (S.) sybleae, S. (S.) thomsenae. 

———. American biting midges of the heleid genus Mono- 
helea. Figures 18,19. Published June 17, 1953 . 

New species: Monohelea (Monohelea) lanet, M. (M.) macfiei, M 


(M.) texana, M. (M.) ornata, M. (M.) stonei, M. (M.) guianae, 
M. (M.) johannseni. 


Vv 


Pages 


357-394 


203-356 


187-201 


439-447 


551-561 


395-438 


57-85 


135-154 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


on — 
GOOLE SHON 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1953 No. 3311 


TWO NEW NAUCORID BUGS OF THE GENUS AMBRYSUS 
By Ira La Rivers’! 


The following new Ambrysi were discovered after the manuscripts 
dealing with United States and Mexican species had been completed, 
and this paper is offered as a supplement to these treatments. Keys 
are included to allow the incorporation of the new species into the 
overall keys for the two regions. 


Family NAucorIDAE 
Genus Ambrysus Stal, 1862 
Ambrysus thermarum, new species 
Figure 1, 0, c 


Description.—A small, pale species, nearly approaching Ambrysus 
funebris La Rivers, 1949, as the smallest species in the genus, and 
greatly resembling the latter in general conspectus; size 7.0-8.0 mm. 
long and 4.5-5.0 mm. wide. Dorsum bicolored, the scutellum and an- 
terior portion yellowish, the posterior area (hemelytra) brownish. 
Venter brownish, contrasting strongly with the yellow-white ap- 
pendages. 

Head glistening, yellow, slightly roughened and weakly punctate, 
protuberant before eyes and with the faintest suggestion of trunca- 
tion in front. Eyes blackish, slightly raised above general head sur- 


* Department of Biology, University of Nevada, Reno, Nev. 
22842252 1 


9) PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


face when viewed posteriorly. Head ratios are: Total length to width 
(including eyes) 19:28 (68 percent) ; anterior distance between eyes to 
posterior distance 12:17 (71 percent) ; anterior distance between eyes 
to inner eye-margin 12:11; posterior distance between eyes to greatest 
length of head posterior to this line 17: 6 (35 percent). 

Pronotum glistening, densely, shallowly punctate; color yellow to 
amber; lateral edges fiaeaie rounded, nonserrate, occasionally with 
some weak pilosity. Curvature 14 percent (average 25:4), postero- 
lateral angles well rounded. Dorsal ratios are: Width between an- 
terior angles to greatest pronotal width 29:51 (57 percent) ; median 
length to greatest width 17:51 (83 percent) ; width between anterior 
angles to distance between anterior angle and posterior baseline of 
pronotum 29:25 (86 percent). 

Scutellum yellow-amber, shiny; shagreened with dense, shallow, 
white-rimmed punctation; ratio of three sides 30: 24: 24. 

Hemelytra uniformly brownish, yellowing medially and in em- 
bolar region; shagreened as is scutellum. Embolium about normal 
for the genus, length-to-width ratio 29:9 (31 percent), rarely with 
some sparse, occasional marginal pilosity. Hemelytra moderately 
exposing lateral connexival edges, which latter have some prominent, 
short yellow pilosity. Hemelytra just attaining abdominal apex. 

Venter reddish brown, contrasting with the light-colored legs. 
Connexival posterolateral angles weakly acute-angulate, gradually 
enlarging posteriorly, not in the least spinose; connexival edges 
smooth, nonserrate, moderately pilose. Female subgenital plate 
apex simply concave, the concavity somewhat angular at its deepest 
point. Male genital process sharply right-angulate, tapering to apex, 
appearing much like a very thin foot, curving abruptly outward. 

Leg I yellow-white, darkening to light amber on tibia-tarsus; 
ratio of length to greatest width of femur 27:16 (59 percent) ; com- 
bined tibia-tarsus, when closed, just attaining, or slightly surpassing, 
adjacent (proximal) end of femur. 

Leg II yellow; ratio of femoral length to median width of ventral 
surface 25:5 (20 percent), length 1.9 mm.; tibia with distal end 
ventrally with one complete transverse terminal row of reddish spines 
set solidly across tip (remnants of the second, subterminal row, when 
complete and unbroken by use, extend about two-thirds of the width of 
tibia), ratio of length to width 22:2 (10 percent), length 1.75 mm. 

Leg III yellow; ratio of length to femoral width 30:5.5 (18 per- 
cent), length 2.1 mm.; distal end of tibia ventrally with the same type 
of transverse spination as in mesotibia; ratio of length to width 37:2 
(5 percent), length 3.0 mm. 

Tibia has the characteristic spinate appearance in contrast to the 
smoother micro-armed femora, with the long, silky, swimming hairs 
on metatibia contrasting with the less hiveute appearance of the 


TWO NEW NAUCORID BUGS—LA RIVERS 3 


mesotibia. All tarsi, except protarsi, 3-segmented; the first seg- 
ment minute; the last two long and slender, terminating in two mod- 
erately curved claws. 

Type and paratypes——USNM 60987. 

Type locality—New Mexico, Sierra County, Hot Springs. 

Material examined —Holotype (male), allotype, and four para- 
types, collected by Loew, September 11, 1874 (P. R. Uhler collection), 
in U.S. National Museum. 

Remarks.—This unique little species, discovered during the course 
of checking the Ambrysi in the National collection, is probably an 
offshoot, as far as present material can indicate, of Ambrysus mormon 
stock. The female subgenital plate is mormonoid in structure, and 
the variable mormon-type male process could easily have given rise 


— 
‘a \ 


Figure 1.—Diagnostic characteristics of Ambrysus bispinus and A. thermarum: a, Apex 
of subgenital plate of 4. bispinus, female holotype (visible only in ventral view), X22; 
b, apex of subgenital plate of 4. thermarum, female allotype, 22; c, genital process on 
caudal edge of tergite V of A. thermarum, male holotype, X22; d, bispinate condition 
of left connexivum of abdominal sternite V of 4. bispinus, female holotype, showing (1) 
anterior posterolateral angle and (2) posterior laterocaudal angle (ventral view, with 
caudal tip of abdomen oriented at the top of the field), 22. 


to the distinctive male process of A. thermarum, which has evolved 
toward the A. woodburyi type to such an extent that, with only the 
male to examine, it would have been considered an aberrant A. wood- 
buryi. At present, it is the most definitely adapted thermal ecad of 
the A. mormon type, but as nothing is known of the structure of the 
type locality, it is impossible to say whether this seems to be due to 
complete isolation or not. 


4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


The following modification of my key to United States Ambrysi 
(La Rivers, 1951) will satisfactorily compare A. thermarwm with its 
closest relatives: 

8 (7). Female subgenital plate simply, broadly, and smoothly concave apically, 
the lateral bordering angles sharp; concavity, measured in terms of 
depth of concavity against length between bordering lateral angles, never 
less than 20 percent ; in clean material embolium nearly unicolorous, only 
very faintly lighter exteriorly, no contrasting colors present. 

buenoi Usinger, 1946 
Female subgenital plate, if simply concave, has (a) rounded lateral angles, 
or (b) if lateral angles are sharp, then depth of concavity is considerably 
less than 20 percent; in clean material embolium always sharply and 
contrastingly transversely bicolored, anterior two-thirds light yellow, 
posterior one-third blackish-brown=— 2-2-2 eee 8A 
8A (8). Male genital process narrowing to apex, strongly, abruptly and almost 
right-angularly turned outward, giving the appearance of a long, 
attenuated foot with a rather sharp heel; female subgenital plate 
simply concave at apex, somewhat angularly so. 
thermarum, new species 
Male genital process not definitely narrowing to apex, and not foot- or 
boot-shaped, although curving variably outward; female subgenital 
plate simply and roundly concave at apex__ mormon Montandon, 1909 


Ambrysus bispinus, new species 


Figure 1, a, d@ 


Description.—A large, darkly mottled species, well rounded and con- 
vex; size 12.0-14.0 mm. long and 7.0-9.0 mm. wide. Dorsum not 
distinctly bicolored (i. e., anterior portion lighter than posterior area), 
more or less uniformly mottled. Venter yellow or brownish, uni- 
colorous and lighter than dorsum. 

Head glistening, yellow-brown, smooth, weakly and vaguely punc- 
tate, slightly and broadly protuberant, and truncate before the eyes. 
Eyes blackish, nearly flush with plane surface of head when viewed. 
posteriorly. Head ratios are: Total length to width (including eyes) 
34:54 (63 percent) ; anterior distance between eyes to posterior dis- 
tance 26:33 (79 percent) ; anterior distance between eyes to inner eye- 
margin 26:25; posterior distance between eyes to greatest length of 
head posterior to this line, 83:8 (22 percent). 

Pronotum glistening, yellowish with brown mottling, sparsely 
punctulate, rugulose behind head; lateral edges smooth, nonserrate, 
rounded, on one specimen with traces of faint pilosity which appears 
to have been rubbed off. Curvature 10 percent (average 52:5); 
posterolateral angles well rounded. Dorsal ratios are: Width be: 
tween anterior angles to greatest pronotal width 53:73 (73 percent) ; 
median length to greatest width 39:73 (53 percent) ; width between 
anterior angles to distance between anterior angle and posterior base- 
line of pronotum 53: 52. 


TWO NEW NAUCORID BUGS—LA RIVERS a 


Scutellum glistening, yellow medially and on posterior angle, 
yellow along lateral edges, reddish brown elsewhere; shagreened with 
dense, coarse, white-spotted punctures; ratio of three sides 70: 50: 50. 

Hemelytra mottled brown and yellow, glistening; shagreened as is 
scutellum. Embolium slightly narrow for the genus, length-to-width 
ratio 60:18 (30 percent), with no detectable marginal pilosity in the 
specimens examined. Hemelytra moderately exposing connexival 
margins, which latter have sparse, short pilosity, concentrated some- 
what at the posterolateral angles. Hemelytra just, to not quite, at- 
taining abdominal] tip. 

Venter yellow to brown, the legs usually not markedly contrasting 
with the body in color. Connexival posterolateral angles acute-ang- 
ulate, short, gradually increasing in size posteriorly, not in the least 
spinose ; margins with some weak dentation, detectable only with con- 
siderable magnification, on all segments except I and II. Female 
segment V (the insect being oriented so that the abdominal tip is at 
the top of the field) possesses two posterolateral connexival processes ; 
the exterior posterolateral angle and an interior, prominent, blacker, 
rather blunt elongation of the caudal margin of segment V lying 
adjacent to the posterolateral angle. I have referred to this in other 
species as the laterocaudal connexival angle. Tip of female sub- 
genital plate strongly and distinctively quadrisinuate, the lateral ter- 
minal angles long and rather sharp, but not extending as far caudad 
as the median, paired sinuosities which lie close together as a doubly 
rounded, blunt process longer than the lateral spines. Character of 
the male genital process unknown. 

Leg I amber; ratio of length to greatest femoral width 48:31 
(65 percent) ; combined tibia-tarsus, when closed, just attaining adja- 
cent (proximal) end of femur. 

Leg II amber; ratio of length to median width of femur 46:10 
(22 percent), length 3.4 mm.; tibia with distal end ventrally with 
one-and-one-half transverse rows of reddish spines, the terminal row 
complete, the subterminal row, as indicated above, only half complete; 
ratio of length to width 40 :4.5 (11 percent), length 3 mm. 

Leg ITI amber; ratio of length to median femoral width 57:11 (19 
percent), length 4.2 mm.; tibia with same type of transverse terminal 
spination as in mesotibia, ratio of length to width 70:5 (7 percent), 
length 5.2 mm. 

Tibia possesses the typical spinate appearance in contrast to the 
smooth femora; long, golden, silky, swimming hairs are present on the 
metatibia and lacking on the mesotibia. Al tarsi, except protarsi, are 
3-segmented, the first segment minute, the last two long and slender, 
terminating in two moderately curved claws. 

Type and paratype-—USNM 60988. 


6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Type locality.—México, Veracruz, Jalapa. 

Material ecamined.—Holotype (female) and a single female para- 
type from México, Oaxaca, Camotlan, collected by W. S. Miller, April 
14, 1944, in the U. S. National Museum. 

Remarks.—In external appearance, A. bispinus is superficially in- 
distinguishable from A. cosmius and, as such, has probably shared 
the latter’s confusion as A. stgnoreti Stal, 1862, in collections. How- 
ever, the structure of the female subgenital-plate apices is quite dif- 
ferent in the two species. In this respect, A. bispinus most closely 
resembles A. guttatipennis Stal, 1876, but while the subgenital plates 
might not be able, alone, to separate all variables of both species, the 
possession of primary and secondary connexival angles on abdominal 
segment V, with present material, sharply sets A. bispinus off from 
all other Ambrysi of the signoreti group. ‘These two angles are more 
specifically referred to as the posterolateral and laterocaudal con- 
nexival angles, and the laterocaudal is lacking in all other known 
members of the group. It has shown up in other, distant sections 
of the genus, being known in A. parviceps Montandon, 1897, A. hun- 
gerfordi Usinger, 1946 (both in the pudicus group), and A. puncti- 
collis Stal, 1876 (a monotypic group). It is this character upon 
which the specific name of A. bispinus is based. In addition, the 
male genital process, when known, will furnish more evidence of the 
species’ affiliations, either supplementing or diverting present indica- 
tions. . 

The following couplets will enable the new species to be incorpo- 
rated in my key to Mexican Ambrysi now in press: 

24 (15). Female subgenital plate, viewed ventrally, asymmetrical on left side, 
where a prominent flap occupies the border beyond (laterad of) the 
left lateral angle; right border scarcely produced. 

dilatus Montandon, 1910 
Female subgenital plate, viewed ventrally, not markedly or noticeably 
asymmetrical from one lateral border to the other_____-________ 25 

25 (24). Female possessing both primary and secondary posterolateral connexi- 
val spines on abdominal sternite V (i. e., the primary angle being the 
posterolateral angle proper, the secondary angle being the laterocaudal 
ANSIC) oo 32 wy i ee bispinus, new species 


Female lacking the secondary (laterocaudal) angle. 
remaining Mexican species of the signoreti group 


TWO NEW NAUCORID BUGS—LA RIVERS 7 


References 
La Rivers, Ira 
1949. A new species of Ambrysus from Death Valley, with notes on the 
genus in the United States (Hemiptera: Naucoridae). Bull. 
Southern California Acad. Sci., vol. 47 (1948), No. 3, pp. 103-110. 
1950. The meeting point of Ambrysus and Pelocoris in Nevada. Pan-Pacific 
Ent. vol. 26, No. 1, pp. 19-21. 
1951. <A revision of the genus Ambrysus in the United States (Hemiptera: 
Naucoridae). Univ. California Publ. Ent., vol. 8, No. 7, pp. 277- 
338, illustr. 
— The Ambrysus of Mexico (Hemiptera: Naucoridae). Univ. Kansas 
Sci. Bull. (In press.) 
MonTANnpon, A. L. 
1897. Hemiptera cryptocerata.—Sous-fam. Cryptocricinae. Verh. zo6l.-bot. 
Ges. Wien, vol. 47, pp. 6-23. 
1909. Naucoridae. Descriptions d’especes nouvelles. Bull. Soc. Roum. Sci., 
Bucharest, vol. 18, No. 1, pp. 43-61. 
STA, CARL. 
1876. Hnumeratio Hemipterorum. Svenska Vetensk. Akad., vol. 14, pt. 5, 
No. 4. 
USINGER, ROBERT L. 
1946. Notes and descriptions of Ambrysus St4l with an account of the life 
history of Ambrysus mormon Montd. (Hemiptera: Naucoridae). 
Bull. Univ. Kansas, Sci. Bull., vol. 31, No. 1, pp. 185-210. 


U, 8. GOVERNMENT PRINTING OFFICE: 1953 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


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SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1953 No. 3312 
TWO NEW SCALE-MITE PARASITES OF LIZARDS 


By R. F. Lawrence! 


Dr. G. W. Wharton, of Duke University, Durham, N. C., has kindly 
submitted to me for identification a number of scale mites from lizards 
of the families Gekkonidae and Iguanidae. The material, from the 
Philippine Islands, consists of one series taken from a lizard at Fort 
McKinley, Rizal, and another from the gecko Hemidactylus frenatus 
in a house at Manila. Both series consist of two quite different species 
of Geckobia. These are described in this paper as new and both of 
them have been taken at the two localities mentioned above. There 
is thus good reason for assuming that the hosts of the two series are 
the same and that the “lizard” of Fort McKinley is identical with 
the gecko H. frenatus from which scale mites were taken at Manila. 
All these parasites were collected by Dr. H. L. Keegan, Third General 
Medical Laboratory, at Fort McKinley, Rizal, Philippine Islands. 

A further series, consisting of a large number of adults and larvae, 
were collected from the iguanid lizard Sceloporus undulatus, at Dur- 
ham, N. C., by Dr. A. S. Pearse; these have been identified as Banks’ 
New World pterygosomid Geckobiella texana. 

Several species of the scale mite parasite Geckobia may live on the 
same host; Hirst (1925, p. 173) has already pointed this out, saying, 
“When more than one form of Geckobia is found on the same host, 
one lives beneath the ventral scales and is flattened, being considerably 
wider than long and having scales instead of hairs on the venter. 
The second form occurring on the same host is usually to be discovered 
between the claw and pad of the toes, between the laminae of the pad, 
or between the toes themselves; this form is practically spherical in 
shape and has hairs instead of scales on the venter.” Although there 


1 Natal Museum, Pietermaritzburg, Union of South Africa, 
228383—53 9 


10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


is no definitive information that the two strikingly different species 
of Geckobia here described actually illustrate Hirst’s observation, it 
will probably be found that the two forms live on different parts of 
the body of the same host. 
Family PrERYGOSOMIDAE 
Genus Geckobia Mégnin, 1878 
Geckobia keegani, new species 
FIGURES 2, 3 

Material examined—Three females, cotypes, from a lizard, Fort 


McKinley, Rizal, Philippine Islands, collected by H. L. Keegan, Octo- 
ber 1948, USNM 1931; also nine females bearing same locality data 


Figure 2.—Geckobia keegant, new species, dorsal surface. 


SCALE-MITE PARASITES—LAWRENCE 11 


b C d 


Ficure 3.—Geckobia keeganit, new species: a, Dorsal scute; b, peripheral dorsal hair; ¢, 
anterior ventral hair; d, posterior ventral hair. 


as the cotypes, and five females from Hemidactylus frenatus, from 
Manila, Philippine Islands. 

Body almost circular; dorsal surface as in figure 2, with fairly 
long, almost parallel-sided hairs differing very little in length, but 
posteriormost slightly longer than anteriormost. Dorsal scute well 
defined, with about 20 hairs differing very little in general appear- 
ance and size from the remaining dorsal hairs, though a little shorter 
than the longest of these. 

Ventral surface with a density of hairs similar to that of the dorsal 
surface but reaching further back, almost to the anal field; the dis- 


12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


crepancy in the length of these hairs far greater than in those of the 
dorsal surface, anteriormost a little less than half as long as posterior- 
most, latter a little shorter than the longest dorsal hairs. 

Mouth parts normal, pedipalps short and stout, both second and third 
segments above with a setiform hair, that of the second weakly barbed 
and only a little stouter than that of the third, which is smooth. 

Legs equal-sized, rather small in proportion to the body. Pos- 
terior common coxa differing from most species of Geckobia in having 
two spurlike hairs on its anterior and only two, one, or even none 
on its posterior margin; usually only three or four hairs, instead of 
the normal five, are found on this coxa and these are comparatively 
small and weak. The basal segments of the legs without spurlike 
hairs ventrally; but the second segment in all legs with a rather 
long, slender, and weakly barbed hair above. 

Dimensions.—Width of body 0.603 mm., length (including mouth 
parts), 0.632 mm. 

Remarks.—The species closely resembles Geckobia simplex Hirst, 
1926, described from Hemidactylus leschenaulti, Madras, India, in 
the shape of the hairs of the dorsal surface and in the form of the 
dorsal scute. It differs from this species in having only about half 
the number of hairs on the dorsal scute, these being also more similar 
to the remaining body hairs in length and thickness than is the case 
in G. simplex. It further differs in there being very little difference 
in the shape of the dorsal hairs of the second and third palpal 
segments. 


Geckobia philippinensis, new species 
FIGURES 4, 5 


Material examined—Two females, cotypes, from a lizard, Fort 
McKinley, Rizal, Philippine Islands, collected by H. L. Keegan, 
October 1948, USNM 1982. 

Body much wider than long, dorsal surface as in figure 4, with 9 or 
10 large, wide hairs in the area usually occupied by the dorsal scute, 
those in the middle of the body much smaller, while towards the pe- 
riphery posteriorly and laterally, the hairs become progressively more 
elongate and pointed. A single pair of centrally situated hairs much 
longer than the predominantly short hairs by which they are sur- 
rounded. Dorsal scute not defined. 

Ventral surface as in figure 5, a, which represents a section in the 
middle line of the body extending from its anterior to its posterior 
margin; the transition from the small type of cylindrical hair to 
scales of lanceolate form, sudden and without intermediate types of 
hair; the scales toward the posterior periphery becoming progres- 
sively more elongate and narrow. 


13 


SCALE-MITE PARASITES—LAWRENCE 


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14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Mouth parts normal, pedipalps short, with rounded segments, sec- 
ond segment above with a stout plumose hair several times thicker 
than that of the third segment, which is smooth and setiform. 

Leg IV distinctly longer and a little stouter than the remaining 
ones; posterior common coxa as in figure 5, e, with five very large 
and distinct conical spurlike hairs; all legs with a fairly stout 


Figure 5.—Geckobia philippinensis, new species: a, Hairs of ventral surface in midline of 
body; 6, peripheral dorsal hair; c, small hair from middle of dorsum,‘d, a scalelike ven- 
tral hair; e, posterior common coxa. 


plumose hair on the ventral surface of segments 2 and 3 and a 
long setose hair on the dorsal surface of segment 3, that of leg 
IV weakly plumose. 

Additional material evamined.—Two females from the same host 
and locality as the types. Dr. Wharton sent with his material the 
lizard host from which the Manila specimens were taken. From the 
body scales of this gecko, Hemidactylus frenatus, I obtained a single 
female of the species described above. 


SCALE-MITE PARASITES—LAWRENCE 15 


Dimensions.—Width of body 0.400 mm., length (including mouth 
parts), 0.287 mm. 

According to Hirst’s key (1925, p. 174), this species would be brack- 
eted with Geckobia australis Hirst, 1926, but in width of body and 
general form of dorsal hairs and ventral scales, it seems to resemble 
more closely G. hindustanica Hirst, 1926, described from Hemidacty- 
lus leschenaulti, Madras, India. It differs from G. hindustanica in 
the complete absence of a dorsal scute and in the relatively larger size 
of leg IV, as well as in other details. 


Genus Geckobiella Hirst, 1917 


This genus was erected by Hirst (1917, p. 188) for the scale mites 
of Iguanid lizards, and thus far its members have been found to 
occur only on species of the genus Sceloporus. It would be of great 
interest to know what types of scale mites occur on the other numerous 
genera of the family Iguanidae in the New World. 

The eyes, which are not mentioned in the description of the genus, 
consist of a single pair situated in much the same position as those 
of Geckobia and Zonurobia, Lawrence, 1935, being located anteriorly 
near the lateral margin of the body. They are small, only a little 
larger han the ringlike sockets of the dorsal hairs (fig. 6, 6). 


Geckobiella texana (Banks), 1905 


Fiaures 6, 7 


Geckobia texana Banks, Proc. Ent. Soc. Washington, vol. 8, p. 134, 1905. 
Geckobiella terana (Banks), Hirst, Ann. Mag. Nat. Hist., ser. 8, vol. 19, p. 
138, 1917. 


Material examined—Numerous adults and larvae of this species 
collected by A. S. Pearse from Sceloporus undulatus at Durham, 
N. C., in January 1950. 

Adult female—Hirst (1925, p. 200, fig. 19) figured only the ventral 
surface of this species, and the opportunity is now taken to give an 
illustration of the dorsal surface (fig. 6). The dorsal hairs are much 
more numerous in some specimens than in others, there being probably 
a certain amount of variation in this respect. 

The mouth parts are longer and more robust than in most genera 
of Pterygosomidae; the free portion of the peritremes reach to the 
distal end of the second palpal segment; the second palpal segment 
with a fairly long slender hair of equal thickness throughout, not 
pointed at the apex, with fine barbs; dorsal hair of the third palpal 
segment longer, pointed apically and smooth or almost so; fourth pal- 
pal segment with a single seta above, claw stout, conical, and somewhat 
curved. 


16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


a x a 
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Ficure 6.—Geckobiella texana (Banks): a, Dorsal surface of female; b, eye. 


Leg I only a little longer than IV, distinctly stouter than the re- 
maining legs; segments 2 and 3 of all legs with a setiform hair above, 
finely barbed. 

Adult male—The single specimen which appears to be an adult 
male is considerably smaller than the female and with far fewer hairs 


SCALE-MITE PARASITES—LAWRENCE 17 


Ficure 7.—Geckobiella texana (Banks): a, Dorsal surface of larva; b, penis (?). 


on the dorsal surface; otherwise it does not differ from the female; 
the chitinous genital structures are rather ill-defined, the large penis 
(%) (fig. 7,5), projecting slightly from the genital aperture at the 
posterior extremity of the body. 

Larva—Dorsal surface with very few symmetrically disposed hairs, 
as in figure 7,a. Eyes present in the same position as the adult. Ven- 
tral surface entirely without hairs or setae except for a single seta at 
the base of legs I and III. Mouth parts closely resembling those of 
the adult, but the peritremes much shorter, reaching to a little below 


18 PROCEEDINGS OF THE NATIONAL MUSEUM YOL. 103 


the base of the second palpal segment, and the claw of the palp more 
slender. Legs shorter than in the adult. 

Dimensions —Female : Length of body 0.9138 mm.; width, 0.487 mm. 
Male: Length of body 0.652 mm.; width, 0.326 mm. Larva: Length 
of body 0.740 mm. ; width, 0.413 mm. 


References 


Hirst, ARTHUR STANLEY 
1917. On some new mites of the suborder Prostigmata living on lizards. 
Ann, Mag. Nat. Hist., ser. 8, vol. 19, p. 136. 
1925. On parasitic mites of the suborder Prostigmata (Trombidioidea), on 
lizards. Journ. Linnaean Soc. London, vol. 36, p. 173. 
LAWRENCE, R. F. 
1935. The prostigmatic mites of South African lizards. Parasitology, vol. 27, 
No. 1, p. 6. 


U.S. GOVERNMENT PRINTING OFFICE: 1953 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


issued 1% 
PER 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington : 1953 No. 3313 


NOTES ON THE BIOLOGY AND IMMATURE STAGES OF A 
CRICKET PARASITE OF THE GENUS RHOPALOSOMA 


By AsHiry B. Gurney’ 


Introduction 


For more than three-quarters of a century wasps of the genus 
Rhopalosoma Cresson, 1865, have attracted the interest of hyme- 
nopterists because characters of the adult have left doubt regarding 
family relationships. Although the family Rhopalosomatidae? was 
proposed for the genus in 1896, there remains uncertainty as to what 
other genera are to be included in the family.’ Neither is it clear 
which of the related families is most closely allied to the Rhopalosoma- 
tidae. Biological studies of Rhopalosoma together with a critical 
examination of the immature stages may be helpful in clarifying these 
matters. Until now the only original published information concern- 
ing the biology and immature stages is that by Hood (1914). He col- 
lected a larva which was reared to maturity and identified as R. poeyi 
Cresson, which is currently referred to R. nearcticum Brues. Hood’s 
account is brief and, while it is a good general indication of the 
biology, contains no detailed morphological descriptions. The recent 
capture of a larva of Rhopalosoma at Falls Church, Va., has given 


1 Bureau of Entomology and Plant Quarantine, Agricultural Research Administration, 
U. S. Department of Agriculture. 

* Originally given as Rhopalosomidae by Ashmead (1896). An adult specimen of the 
genus has been illustrated by Westwood (1874, pl. 24, fig. 9). The species shown there is 
Rhopalosoma aenigmaticum (Westwood) of Hispaniola. 

® Readers are referred to Brues and Melander (1932, p. 499), Brues (1943), and Krom- 
bein (1951). 


228353—53 19 


20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


impetus to the gathering of biological data, and the material at my 
disposal has also permitted morphological studies to be made on 
several of the immature stages. 

I am much indebted to Dr. Henry K. Townes, of the North Carolina 
State College (in absentia), for lending specimens of Rhopalosoma 
and for supplying collecting records and giving helpful suggestions. 
My colleagues Dr. B. D. Burks, K. V. Krombein, and Dr. W. H. 
Anderson have also advised me generously. David G. Shappirio, of 
Harvard University, and Dr. T. H. Hubbell, of the University of 
Michigan, have kindly supplied data on parasitized crickets preserved 
at the University of Michigan Museum of Zoology. 


Biological observations 


The larva found at Falls Church ‘ was in the fourth instar, attached 
to the left side of an adult female of Hapithus agitator Uhler. While 
I was beating deciduous underbrush for psocids, on September 30, 
1950, the cricket was knocked into a collecting umbrella. The left 
hind leg was held at an unnatural angle with the body and the cricket 
was rather inactive and easily picked up. Leaves and other organic 
material were added to the jar in which the insect was placed, but the 
cricket was not seen to feed. The next day it scarcely changed its 
position, though the antennae occasionally moved. Meanwhile, the 
ventral surface of the parasite remained securely attached to the 
cricket and no movement was noticed. A 7 p.m. on the evening of 
October 2 the larva was beginning to shed its skin, as shown by the 
appearance of the white integument of the fifth instar where the 
fourth instar skin had split lengthwise along the middle of the 
dorsum. The cricket had then lost its foothold and was dying on its 
side, though its antennae still moved. 

By 8:30 p.m. the larva (which was entirely white except for traces 
of pale brown about the mandible) had left the dead cricket and had 
moved an inch or soaway. ‘The larva was placed in a pill box of soil 
where it gradually darkened to a creamy yellow color. On the day 
following the molt the larva made frequent twisting contractions as 
if trying to burrow. The head was directed downward most of the 
time and one silk strand coming from the mouth was noticed. The 
larva did not succeed in becoming completely covered with soil; dur- 
ing the next day it was relatively inactive, with the head uppermost 


4 Generic identification of the parasite is primarily based on the very close agreement 
with the situation reported by Hood (1914). It may eventually be shown that more than 
one nominal form of Rhopalosoma inhabits the United States and that some specimens 
from the vicinity of Washington, D. C., are distinct from R. nearcticum, the type locality 
of which is Kissimee, Fla. 


CRICKET PARASITE—GURNEY Pi 


and showing occasional movements of the mouth parts. By the morn- 
ing of October 5, it was decidedly quiescent and evidently dying, and 
by noon it had died and so was preserved in alcohol (pl. 1). If 
several inches of fine sand had been provided instead of heavy gar- 
den loam, perhaps the insect would have entered the soil and suc- 
cessfully spun a cocoon. 

The present larva was attached to the cricket on the membrane 
between the terga and sterna of abdominal segments 1 to 5. After 
the fifth instar had left, the cricket (pl. 1) bore three exuviae 
visible from above, representing the second, third, and fourth instars. 
In each the head was directed posteriorly and located farther posteri- 
orly on the cricket than that of the previous instar. The strongest 
single point of larval attachment was apparently just anterior to the 
parasite’s modified anus; there the attachment was quite secure. The 
larval body was noticeably curled, but the entire ventral surface was 
closely appressed to the cricket. The exuviae of larval instar 1 was 
situated transversely on the ventral surface of the cricket, in a groove 
just behind the left hind coxa. The head end of the skin was directed 
mesally on the cricket and extended almost to the sternal plate of the 
first abdominal segment, while the posterior end was beneath the ends 
of succeeding exuviae. Remnants of the eggshell occurred with the 
exuviae of the first instar. On the body of the cricket, in part be- 
neath the cast skins and also directly adjacent to them and extending 
briefly onto the left tegmen (front wing), was an accumulation of 
dried mucus or similar material. This was scalelike in appearance 
and I first mistook it for exuviae. The larval head was not inserted 
through the cricket’s body wall, nor was there any definite opening, 
but the membrane where the last feeding mandibles (fourth instar) 
were applied showed signs of having been pierced. The points of 
contact for the mandibles of the second and third instars were covered 
with the exuviae of later instars. 

A second larva of Rhopalosoma in the U.S. National Museum was 
collected on a last instar male nymph of Hapithus sp. at Marietta, Ga., 
August 9, 1947. The larva is in the fifth instar, though it probably 
was captured in an earlier stage since the host cricket is also preserved. 
The exuviae which were attached to the left side of the cricket agree 
with those of the Falls Church specimen so far as has been noted. 

The third larva of Rhopalosoma examined (fig. 9, e, f) was preserved 
in the fourth instar after having been collected on an adult female of 
Hapithus agitator at Fuguay, N. C., September 12, 1950, by Townes, 
Rabb, and Howden (North Carolina State College collection) (pl. 1). 
The head of the final feeding stage of the larva was on the cricket’s 
fifth abdominal segment, with the mandibles applied to the mem- 
brane close to the spiracle and nearer the tergum than to the 


22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


sternum. <A crust of a dried secretion was present, as in the case of 
the Falls Church larva. Though a crust may have been associated 
with the Marietta, Ga., material, none was seen. The eggshell and 
first larval exuviae of each of the three larvae occurred immediately 
behind the hind coxa of the cricket. Likewise, the hind leg of each 
cricket, adjacent to the parasite, was held outward as shown in the 
photographs. 

A fourth larva examined is one of several which Dr. Townes has 
told me of finding on Hapithus and Orocharis. Of this particular 
larva he has written, “On September 12, 1950, I collected (at Fuguay, 
N. C.) a large Orocharis nymph with a one-third grown larva on it. 
I kept the cricket in a jar and its parasite prospered. On September 
17 I found the remains of the demolished nymph in the jar and the 
Rhopalosoma larva, now creamy white with a large head and pro- 
jecting mandibles, busily hitching itself along with its mandibles. 
It was very restless, so I put it in some damp sand. The larva 
promptly worked into the sand and came to rest in one corner of the 
jar, under the sand. Three days later it had completed an elongate, 
castaneous cocoon.” 

The cocoon was kept indoors by Dr. Townes, at a temperature in 
the neighborhood of 55° F. In late February he mailed it to me and 
suggested that dissection to determine whether pupation had occurred 
might be advisable. The parchmentlike cocoon was 14 millimeters 
long, about 5 millimeters in diameter, and rounded at the ends. The 
inner surface was a lighter chestnut color than the outside and scarcely 
any individual threads of silk were apparent except on the outside. 
When the cocoon was opened on February 23, the larva was doubled 
up in the middle of it with about one-fourth of the cocoon’s length 
empty at each end. Pupation had not occurred, nor had there been 
any casting of larval skins or passing of fecal material, but the larva 
appeared perfectly healthy. After preservation it was 6 millimeters 
long. The mandibles bore no teeth. 

The original observations of Hood (1914) were based on a larva 
taken at Plummer’s Island, Md., October 6, 1912. The host, Orocharis 
saltator Uhler, was quite active when found, as it was said to be 
“scampering over the forest floor as rapidly as a large abdominal 
protuberance and a nearly functionless hind leg would permit.” The 
anterior end of the parasite was at the host’s eighth abdominal seg- 
ment. ‘This specimen grew rapidly, left the cricket three days after 
capture and entered the soil, emerging indoors as an adult about 
March 1, 1913. The information Hood gave concerning the exuviae 
and the quiescent state of the host just prior to the parasite’s final 
visible molt agrees fully with my findings. The cocoon from which the 
adult emerged has recently been opened and the fifth instar exuviae 


CRICKET PARASITE—GURNEY 25 


were relaxed and studied. The mandibles of the exuviae differ some- 
what in shape from those of the Falls Church larva and they bear no 
teeth. 

In the University of Michigan Museum of Zoology are at least 20 
pinned cricket specimens bearing parasitic larvae presumed to be of 
one or more species of Rhopalosoma. I have not examined them, 
but detailed host data have been furnished me, together with the state- 
ment that the larvae vary considerably in size and that all are brown. 
The specimens were collected by Dr. T. H. Hubbell and several asso- 
ciated collectors and former students. The hosts, with the number 
of parasitized specimens, States where collected, and generalized 
dates of collection are as follows: Hapithus agitator agitator Uhler, 
seven specimens, southeastern Indiana, central Missouri, Mississippi, 
Tennessee, all collected in the last half of August; H. agitator quad- 
ratus Scudder, four specimens, Florida, late July to early September ; 
H. brevipennis Saussure, four specimens, Florida, late July to mid- 
November ; Orocharis saltator Uhler, four specimens, July 4 to July 
29. The host associated with an additional larva from Florida is 
not recorded. These records represent the parasitism of this type 
occurring among several hundreds of Hapithus and Orocharis 
specimens. 

The specimen of Hapithus agitator on which the Falls Church larva 
was parasitic was preserved in alcohol and later dissected to determine 
the condition of its internal organs. Scarcely any traces of food 
were in the digestive tract, none of recent ingestion. Reproductive 
organs were evidently intact, but there were no well-developed eggs 
or other signs of normal functioning of the organs. 

The impression created was that the cricket had been weakened 
tc a point where the drain on its vitality finally caused death, though 
there had been no mechanical destruction of organs. 

Various detuils of the biology remain to be clarified, but the main 
outline seems evident. Upon hatching, the larva probably crawls part 
way from the eggshell and starts its development in a more or less 
transverse position. At the first molt the larva apparently shifts 
the anterior part of the body so that it extends longitudinally along 
the side of the cricket, though keeping the same place of anal attach- 
ment. The third and fourth instars each move the head posteriorly 
on the cricket at molting time. Sharp mandibles occur in the second, 
third, and fourth instars, and these apparently pierce the cricket’s 
integument and permit fluids to be taken into the mouth. Definite 
mouth parts of the first instar have not been seen, but may occur. 
The fifth instar does not feed but is an inactive stage that leaves the 
host and spins a cocoon in the soil. Pupation is evidently delayed, 
perhaps until near emergence time of the adult. 


24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


Regarding habits of Rhopalosoma adults, Dr. Townes has furnished 
the following notes: “I have seen the adults on several occasions and 
usually was fortunate to catch them when seen. I should say that 
in the Upper and Lower Austral Zones of the Atlantic States they 
are widespread and not uncommon in the right habitat. This appears 
(o be places of dense, shrubby vegetation where there is considerable 
humidity, as along: stream bottoms and seashores, or, in other words, 
the same sorts of places where Hapithus and Orocharis abound. The 
adults may be flushed from the bushes. They fly up and alight again 
like an ophiomine ichneumonid. They may be distinguished from 
ophionines in flight only by a slightly stockier appearance. At Long 
Beach, North Carolina, at twilight on July 9, 1949, I saw about 10 
all together in normal flight, cruising in a rather slow but erratic 
manner just over the beach shrubbery, again like an ophionine ich- 
neumonid in its twilight flight. They were flying until no longer 
visible in the gathering dark. I could catch only two of these and 
they were both males.” 

The following collection dates accompanying specimens identified 
as Rhopalosoma nearcticum are available as an indication of when 
adults are active: Maryland: Mayo Beach, September 14, 1947, female 
(Townes) ; Berlin, July 19, 1932, female. North Carolina: Long 
Beach, July 9, 1949, two males (Townes) ; Wallace, August 2, 1949, 
female (Townes). South Carolina: Greenville, September 2, 1940, 
male (H. and M. Townes). Georgia: Stone Mountain, August 18, 
1949, female (Fattig) ; Atlanta, August 11, 1946, female. Florida: 
About a dozen specimens, essentially all reported by Brues (1943), 
ranging from June 9 to August 13, but mostly collected in June and 
July. 

In addition to the foregoing States, nearcticwm has been recorded 
from Missouri and Kentucky by Krombein (1951). 

The known hosts of Rhopalosoma in the United States are Orocharis 
saltator Uhler and Hapithus agitator Uhler. Judging from the rec- 
ords of parasitized crickets preserved in the University of Michigan 
Museum of Zoology, Hapithus brevipennis Saussure appears to be a 
host also. The distribution of O. saltator, as stated by Hebard (1931, 
p. 217), covers the entire southeastern United States, extending north- 
ward to southeastern Pennsylvania, Illinois, and Missouri, and un- 
doubtedly to eastern Texas in the Southwest. In southern Florida a 
primarily West Indian species, Orocharis gryllodes (Pallas), occurs. 
H. agitator occurs as far northeast as Long Island, and otherwise is 
largely comparable to O. saltator in distribution (Blatchley, 1920, 
p. 740; Hebard, 1931, 1938; Rehn and Hebard, 1916, pp. 308-810). 
Two geographic subspecies are recognized, the southeastern one, H. 
agitator quadratus Scudder, living from the Atlantic Coast to Browns- 


CRICKET PARASITE—GURNEY 25 


ville, Tex., and in the East apparently merging with typical H. agita- 
tor in central Georgia and along the “fall line” in North Carolina. H. 
brevipennis inhabits Florida, Georgia, and Louisiana, and doubtless 
other southeastern States. It may be that any of the crickets men- 
tioned above are satisfactory hosts of Rhopalosoma. In southeastern 
Pennsylvania adults of both Orocharis and Hapithus begin to appear 
by early August, and in all of the more northern States the crickets 
pass the winter as eggs, and adults persist until frost time in the fall. 
Blatchley (1920) reported finding hibernating adults of H. agitator 
in Florida and suggested that they represented a fall generation. The 
subfamily Eneopterinae, to which Hapithus and Orocharis belong, is 
primarily tropical, and various genera are well represented in the 
Neotropical Region, from which Brues (1943) has recognized four 
species of Rhopalosoma. 


Morphological descriptions 


Mature larva (fifth instar from Falls Church, Va.).—General shape 
as in plate 1; strongly mandibulate; legs entirely absent; abdomi- 
nal segments moderately well demarked, less so posteriorly, those be- 
yond segment 8 not readily separated; two thoracic and eight 
abdominal pairs of spiracles. 

Head in frontal view as in figure 8, f; occipital border broadly and 
evenly rounded (asymmetrical in the figure because it is bordered by 
uneven cervical folds) ; vertex and frons not individually demarked, 
the integument shiny, slightly irregular, not noticeably punctate; no 
frontal arms evident; paired conspicuous spots (ey), presumed to be 
vestigial eyes, just above base of clypeus; antenna reduced to sensory 
pit with traces of facetlike division, situated well to the side of frons, 
no apparent projecting sensilla; a presumed sensory area represented 
by about nine poorly defined, pigmented spots (sa), with tiny seta at 
lateral margin of group; frons and vertex with very sparse, short 
setae, grouped as illustrated; six strong clypeal setae; labrum con- 
spicuously bilobed, each lobe with two major setae, lateral and an- 
terior margins with microsetae, an irregular row of pits across each 
lobe, some bearing short setae; mandibles heavily sclerotized, apex 
prominent and narrowly rounded, basal third swollen laterally and 
with a single seta, biting margin thin, demarked from main mandib- 
ular structure by groove, and sharply but delicately toothed. (Left 
mandible with four well-spaced teeth along middle third of biting 
margin, followed by closely set group of four; right mandible with 
two teeth, others perhaps lost by breakage.) Maxilla with several 
medium-sized setae as illustrated, apex near tip of mandible sharply 
rounded and with asperities, maxillary palpus barrel shaped, galea 


°6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


equally elongate but of smaller diameter; labium with palpi subequal 
to galeae, area of sericteries with two angular projections posterior to 
weakly sclerotized nonsegmented elongate appendages of uncertain 
homology; apparent opening of silk duct (sd) between bases of latter 
appendages. 

Anterior spiracle on posterior third of prothorax; second spiracle 
barely anterior to groove separating meso- and metathorax; abdominal 
spiracles on segments 1 to 8, slightly more dorsad than thoracic ones, 
situated near anterior margin of the respective segments; each spiracle 
unspecialized so far as visible under 72-power magnification, consist- 
ing of circular disk with central aperture (internal structures not 
studied) ; no setae on thorax or abdomen apparent; no apparent spe- 
cializations for crawling or clinging; vestigial anus transverse, at 
posterior margin of segment 10. 

Cotoration: General color creamy yellow, darker on head (con- 
spicuous black areas on body shown in photograph are necrotic spots 
developed at time of death). Vertex, frons, and clypeus yellowish; 
eye spots, sensory areas, antennae, and pits at bases of setae pale brown- 
ish; a darkened area anterior to sensory area; a somewhat darkened 
indefinite band across base of clypeus connecting bases of mandibles; 
labrum colorless, except for brown setae and pits; mandibles marked 
as illustrated with rich, dark brown; maxillary and labial palpi and 
galeae dark brown; labium white, elongate appendages in area of 
sericteries tinged with yellowish; spiracles yellowish; no special pig- 
mented areas about nonfunctional anus or elsewhere on thorax and 
abdomen. 

Measurements (in millimeters): Length 8 (would actually be 
slightly longer if body not curved); greatest width of body 3.2; 
greatest width of head capsule 1.8. 

Comments: Fifth instar from Marietta, Ga., more robust than 
larva from Falls Church, Va.; setae on frons and vertex smaller, 
scarcely noticeable; teeth on mandibles lacking. Measurements (in 
millimeters): Body length 8.5; greatest width 4; greatest width of 
head capsule 1.6. 

Penultimate stage larva (fourth instar from Fuguay, N. C.) —Gen- 
eral appearance typically hymenopteriform (fig. 9, e, f) ; head capsule 
and mouth parts not so highly developed as in fifth instar; legless; 10 
pairs of spiracles; modified anus apparently specialized as a hold-fast 
organ. 

Head capsule (fig. 8, c) rather masklike in contrast with that of 
fifth instar, integument very rough and irregularly rugose, a conspicu- 
ous tranverse ridge (epistoma) dorsad of labrum; two pairs of blunt 
protuberances on upper part of face, the media pair (mt) broad and 


PROCEEDINGS, VOL, 103 PLATE 1 


U.S, NATIONAL MUSEUM 


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CRICKET PARASITE—GURNEY OF 


FIGURE 8.—The genus Rhopalosoma Oresson, 1865: a, Mouthparts of fourth instar exuviae; 6, external view 
of sixth abdominal spiracle of fourth instar exuviae; c, front view of head, fourth instar alcoholic specimen; 
d, antenna of fourth instar exuviae; e, anal region, fourth instar exuviae; J, front view of head, fifth instar 
alcoholic specimen; (¢ made from Fuguay, N. C., specimen, all others from Falls Church, Va., material), 


28 PROCEEDINGS OF THE NATIONAL MUSEUM vOL, 108 


irregularly rounded, lateral pair (lt) narrow and evenly rounded; an- 
tennal socket (an, fig. 9, e) circular, minute, traces of sensilla scarcely 
visible under 72-power magnification; labrum membranous except 
mesally and at lateral extremities, strongly bilobed, each lobe bearing 
two small setae about equidistant from center and a minute seta near 
anteromesal margin; exposed apical fourth of overlapping mandibles 
triangular, the apex sharply acute, about seven sharp teeth; approxi- 
mate basal third of each mandible exposed near end of epistoma, bear- 
ing a single, minute seta; a long tapering apodeme extending dorso- 
laterally within head capsule; maxillary sclerome (mxs) anterior to 
base of mandible, shaped as illustrated; labial sclerome (labs) an ir- 
regular, incomplete circle. 

Abdomen with 10 apparent segments; spiracles located as in fifth 
instar; entire dorsal area glabrous; venter of prothorax and cervical 
region bearing minute asperities; dorsal and lateral integument ap- 
pearing as if covered with closely appressed scales (see later descrip- 
tion of exuviae) ; eight abdominal segments with ventral, transverse 
groups of small platelets; hold-fast organ at posterior margin of seg- 
ment 10, transverse, weakly crescent-shaped, with short membranous 
flap extending posteriorly. 

CororatTion: General color dark blackish-brown; ventral region 
(unstippled areas on fig. 9, e, /) unpigmented ; a white middorsal line 
(where split occurs at subsequent molt) extending entire length of 
body; head considerably darker than thorax or abdomen; region of 
frons lighter colored than main epicranial region, two darker pig- 
mented spots (ps) anterior to median tubercles, a wide longitudinal 
white stripe on frons, interrupted on anterior portion of vertex and 
resumed as narrow line on occiput; labrum unpigmented except at 
lateral extremities, on a slender transverse basal median sclerite, and 
mesally at the junction of the lobes; mandibles and scleromes of max- 
illae and labium pigmented with brown as illustrated. Abdomen 
with paired unpigmented spots dorsad of spiracles 3 to 8, and in a 
similar position on abdominal segment 9 (which bears no spiracle) ; 
transversely grouped platelets of ventral surface pale brown, also clos- 
ing sclerites of hold-fast organ and immediately adjacent membrane. 

MrAsurEMENtTsS (in millimeters): Length of body 5.7; greatest 
width of body 2.2; greatest width of head capsule 1.4. 

Comments: Slide-mounted preparations of the cast skin of the 
fourth instar from Falls Church, Va., permitted structures to be 
observed in much greater magnification than was possible with the 
preserved alcoholic larva from Fuguay, N. C. While no significant 
differences in these individual fourth stage larvae apparently occur, 
it is important that the features illustrated and described be attributed 
to the correct specimens. 


CRICKET PARASITE—GURNEY 29 


FIGURE 9.—The genus Rhopalosoma Cresson, 1865: a, Section from lateral area of thorax, showing closely 
grouped platelet structure, fourth instar exuviae; b, integumental asperities on ventral surface of pro- 
thorax, fourth instar exuviae; c, d, specialized platelets on ventral surface of fourth and fifth abdominal 
segments, respectively, fourth instar exuviae; e, f, ventral and lateral views, respectively, of fourth instar 
alcoholic specimen; (e, f, from Fuguay, N.C., specimen, others from Falls Church, Va., material). 


an—antenna labs—labial sclerome mxp—maxillary palpus 
ev—closing valves labr—labrum mxs—maxillary sclerome 
ep—epistoma Ip—labial palpus ps—pigment spots 
es—esophagus lt—lateral tubercle sa—sensory area 

ey—eye md—mandible sd—opening of silk gland 
ga—galea mdc—mandibular condyle or sericteries 
hg—hind gut mt—median tubercle tr—trachael ring 


vp—ventral platelets 


30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Exuviae split lengthwise on dorsum, from epistoma to tenth ab- 
dominal segment; presumed antenna consisting of circular socket 
(fig. 8, d@) with three tiny unpigmented circles in center, each bearing 
a central spot of pigment; mouth parts (fig. 8, @) with mandibles 
the most characteristic and best preserved of the parts; mandible thin, 
much flattened, biting margin with large apical tooth and about 10 
acute smaller teeth in a single row along apical third, an emargination 
a little posterior to the teeth, portion of margin overlaid by labrum 
scarcely pigmented, a well-sclerotized and pigmented region at base 
of mesal margin; posterior margins of mandible with a prominent 
black ventral condyle, a small seta near base of long lateral apodeme 
(presumed mesal apodeme for muscle attachment missing in available 
material) ; labrum largely membranous, with irregular transverse 
wrinkles, a constricted pigmented sclerite mesally, about five subapical 
minute pits arranged transversely on each lateral half; a poorly 
defined, somewhat striated organ leading posteriorly to labial sclerome 
from base of labrum, dorsad of remnants of apparent esophagus 
(es); labial and maxillary scleromes showing somewhat different 
shape in flattened aspect than in natural position on alcoholic larva. 

Integument of most of thorax and abdomen smooth, microscopically 
marked into platelets of different shapes on various areas of the 
body, averaging transversely elongate ventrally, irregularly quadrate 
(fig. 9, a) dorsally on thorax and anterior half of abdomen; ventral 
region of neck and prothorax bearing minute asperities of varied 
character (fig. 9,5), some setiform, others in form of sharp rugosities; 
less developed asperities on ventral surface of abdomen; specialized 
platelets on venter of abdominal segments 1 to 8 (poorly developed 
on 1 to 3 and 8, well developed on 4 to 7), of irregular grouping on 4 
and 5 (fig. 9, ¢), others transverse (fig. 9, d@). Spiracles each sur- 
rounded by narrow irregular area of membrane, the external portion 
a circular flattened disk with small central opening; trachea tapering 
to spiracular opening, with internal ring (tr, fig. 8, 6) which may 
function as closing apparatus. Modified anus (fig. 8, ¢) with well- 
sclerotized and pigmented closing valves (cv), integument at lateral 
extremities wrinkled, that posterior and anterior to valves moderately 
pigmented, a short section of membrane on internal surface an appar- 
ent remnant of hind gut (hg). 

Exuviae of the first three instars are very closely appressed to each 
other along the posterior half of their abdominal sterna, so much so 
that they can scarcely be separated without injury to the specimens. 
The closest attachment appears to be a little anterior to the modified 
anal opening (hold-fast organ). The platelets comprising the micro- 
structure of the integument on the lateral and dorsal surfaces of 


CRICKET PARASITE—GURNEY al 


thorax and abdomen, which in the fourth instar vary from tightly 
grouped on the side of the thorax (fig. 9, a) to narrowly separated on 
the abdomen, are much less closely grouped in the third and second 
instars, while they are so distant and reduced in size in the first instar 
as to resemble very widely spaced spiculelike spots of pigment. 
Mandibles of the third and second instars roughly resemble those 
of the fourth, and bear sharp teeth, there being at least six to eight 
in the third and apparently fewer in the second. The epicranium of 
the first instar has about two pairs of rather large areas of darker 
pigment, but no mouth parts have been observed, though they may 
have escaped detection. There are 10 pairs of spiracles in the third 
and second instars, with the pre-spiracular tracheal ring poorly defined. 
Only six pairs of spiracles have been found on the exuviae of the first 
instar, and the posterior one is poorly developed. Each of the three 
early instars is brown, darker on the head. The venter is pale in 
the third and second instars. Approximate lengths of the exuviae of 
the Falls Chuch, Va., larva are as follows (in millimeters) : Fourth 
6.5; third 3.6; second 2.3; first 1.2. Those of the Marietta, Ga., larva 
are almost identical, but the Fuguay, N. C., larva is larger, the three 
eXuviae measuring as follows: Third 4.4; second 2.6; first 1.3. 

Eggs apparently are broadly elongate-oval; fragments are about 
1.4 millimeters long. 

A basic paper on the larval head of parasitic Hymenoptera is that 
of Vance and Smith (1933). Among the more specialized papers, 
those of G. C. Wheeler (1943) and Beirne (1941) have aided me most 
while studying the anatomy of fifth- and fourth-instar heads, re- 
spectively, of Rhopalosoma. The book by Clausen (1940) is a val- 
uable aid toward understanding the biology and morphology of 
related families, and it should be consulted for references to the im- 
portant work of J. C. Bridwell, F. A. Fenton, R. C. L. Perkins, F. X. 
Williams, F. van Emden, and others. 


Summary and discussion 


Among the biological data additional to those brought out by Hood 
(1914) is the fact that Hapithus agitator is an acceptable host for 
Rhopalosoma in the eastern United States. The effect of the parasite 
on the internal organs of the host, unfortunately based on only one 
dissected cricket, has been described. Details concerning the position 
of the parasite egg and various larval stages have been given. Notes 
on adult habits, essentially lacking previously, make it clear that in 
parts of the Southeast specimens of Rhopalosoma are not nearly so 
scarce as has been supposed. Like many another rarity in collections, 
these insects can doubtless be taken in numbers by those familiar with 


32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


their appearance, time of occurrence, and favored habitats. It is note- 
worthy that on a single day, September 12, 1950, Dr. Townes and his 
associates found two larvae, one on Hapithus, the other on Orocharis. 
Specimens belonging to the University of Michigan show that on three 
other occasions, once in Indiana and twice in Florida, a collector has 
found two instances of parasitism in a single day. 

There remain many unanswered questions concerning the biology 
of the species of Rhopalosoma, some of which may require controlled 
cage studies. Stages earlier than the fourth instar are still known 
from exuviae and fragments of eggshell, and first instar specimens 
are especially desirable for morphological study. 

More study material, including specimens from northern and south- 
ern States, may show that more than one form of Rhopalosoma occurs 
in the United States. Adults of nearcticum were distinguished from 
their Neotropical congeners by Brues (1943) on rather minor morpho- 
logical features, and differences correlated with geographic distribu- 
tion might be better represented in larvae than adults. Of the four 
fifth-instar specimens examined, the ones from Georgia and North 
Carolina and the exuviae of Hood’s Maryland specimen lack teeth on 
the mandibles, while the larva from Virginia apparently has a full 
complement of teeth on the left mandible but only a few basal ones on 
the right mandible. It is not known whether occasional breakage of 
teeth occurs while the larvae are burrowing in soil prior to cocoon 
formation. Some breakage of the teeth of the Virginia larva may 
have resulted from my manipulation of the mandibles with needles 
before I realized the existence of teeth. The three fifth-stage larvae 
have been measured for maximum head width and length of mandibles. 
The ratio of head width to length of mandible is 1.68, 1.86, and 2.0 for 
the larvae from Virginia, North Carolina, and Georgia, respectively. 

Brief examination of a few examples of other families has sug- 
gested certain characters which deserve careful attention in subse- 
quent work of a comparative nature. A chrysidid larva (Chrysis) 
has the second thoracic spiracle located behind the groove separating 
meso- and metathorax, while the corresponding spiracle of Ahopalo- 
soma is slightly in front of the groove. Unlike the condition in 
Rhopalosoma, the second spiracle of a tiphiid larva (Costla) is smaller 
than adjacent ones. This tiphid larva has two maxillary appendages, 
and there are two prominent angular projections on the labium in the 
area of the sericteries. As more details of rhopalosomatid life histories 
become available, comparisons with the situation in the Tiphidae, 
such as has been discussed by Clausen, King, and Teranishi (1927), 
will perhaps be instructive. 

A larva of one of the Larridae (Larra analis Fabricius, which at- 
tacks mole crickets) has the head proportionately smaller than 


CRICKET PARASITE—GURNEY 30 


Rhopalosoma, but as in the latter the maxilla and labium have two 
pairs of appendages each. A conspicuous difference is that the man- 
dible of Zarra has a single preapical tooth in the incisor area, and 
basally on a lower lever there are two heavy molar teeth, or “pros- 
thecae.” The mesal pair of elongate labial appendages of Rhopalosoma 
has apparent homalogues in this larrid. 

The mouth parts of those few larvae of Bethylidae and Dryinidae 
that I have seen are much reduced or otherwise specialized. Illustra- 
tions of dryinid larvae on the bodies of leafhoppers, such as those 
of Haupt (1916), show that the parasite larva sometimes remains 
associated with its exuviae in a manner suggestive of Rhopalosoma. 
The genus Harpagocryptus Perkins, based on a species that Perkins 
(1908) described from Queensland, is of interest in this connection 
because the larva was said to form “a sac on the sides of the abdomen 
of small crickets (Trigonidiidae).” Although originally referred by 
Perkins to the Dryinidae, Bischoff (1927, p. 407) called Harpago- 
eryptus a bethylid, and Reid (1939) suggested that it may be a psam- 
mocharid. As with Rhopalosoma, a knowledge of the immature stages 
of Harpagocryptus might prove of great value when adequate com- 
parative studies eventually are possible. 


References 


ASHMEAD, WILLIAM H. 
1896 Rhopalosomidae, a new family of fossorial wasps. Proc. Ent. Soe. 
Washington, vol. 3, pp. 303-309. 
BERNE, BRYAN P. 
1941 <A consideration of the cephalic structures and spiracles of the final 
instar larvae of the Ichneumonidae (Hym.). Trans. Soc. British 
Ent., vol. 7, pp. 123-190, 31 figs. 
BiscuHorr, H. 
1927 Biologie der Hymenopteren. 598 pp., 224 figs. Berlin. 
BLATCHLEY, W. S. 
1920 Orthoptera of Northeastern America. 784 pp., 246 figs. Indianapolis. 
BRvUES, CHARLES T. 
1921 Correlation of taxonomic affinities with food habits in Hymenoptera, 
with special reference to parasitism. Amer, Naturalist, vol. 55, pp. 
134-164. 
1943 The American species of Rkopalosoma. Ann, Ent. Soc. Amer., vol. 36, 
pp. 310-818, 9 figs. 
BRUES, CHARLES T., AND MELANDER, A. L. 
1932 Classification of insects. Bull. Mus. Comp. Zool., vol. 73, pp. 1-672, 
1121 figs. 
CLAUSEN, CurRTIs P. 
1940 Entomophagous insects. 688 pp., 257 figs. 
CLAUSEN, Curtis P., Kina, J. L., AND TERANISHI, CHO 
1927 The parasites of Popillia japonica in Japan and Chosen (Korea), 
and their introduction into the United States. U. S. Dep. Agr. Dep. 
Bull. 1429, 1-55 pp., 1 pl., 35 figs. 


34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


CUSHMAN, R. A. 
1926 Some types of parasitism among the Ichneumonidae. Proc. Bnt. Soc. 
Washington, vol. 28, pp. 29-51, 51 figs. 
Haupt, H. 
1916 Beitrage zur Kenntnis der Cicadinenfeinde. Zeitschr. Wiss. Insekt- 
enb., vol. 12, pp. 217-2238, fig. 13. 
HEBARD, MORGAN 
1931 The Orthoptera of Kansas. Proc. Acad. Nat. Sci. Philadelphia, vol. 83, 
pp. 119-227, 2 figs. 
1938 Where and when to find the Orthoptera of Pennsylvania, with notes 
on the species which in distribution reach nearest this state. Ent. 
News, vol. 49, pp. 97-103. 
Hoop, J. DOUGLAS 
1914 Notes on the life history of Rhopalosoma poeyi Cresson. Proc. Ent. 
Soe. Washington, vol. 15 (1918), pp. 145-147, fig. 
KROMBEIN, K, VY. 
1951 [Rhopalosomatidae] in Muesebeck, et al., Hymenoptera of America 
north of Mexico. U.S. Dep. Agr. Monogr. 2, pp. 773-774. 
PERKINS, R. C. L. 
1908 Some remarkable Australian Hymenoptera. Proc. Hawaiian Ent. 
Soc., vol. 2, pp. 27-385. 
REHN, J. A. G., AND HEBARD, MORGAN 
1916 Studies in the Dermaptera and Orthoptera of the Coastal Plain and 
Piedmont region of the southeastern United States. Proc. Acad. 
Nat. Sci. Philadelphia, vol. 68, pp. 87-314, 3 pls. 
Rep, J. A. 
1939 On the relationship of the hymenopterous genus Olixon and its allies, 
to the Pompilidae (Hym.). Proc. Ent. Soc. London, ser. B, vol. 8, pp. 
95-102, 2 figs. 
VANCE, A. M., AND SMITH, H. D. 
1933 The larval head of parasitic Hymenoptera and nomenclature of its 
parts. Ann. Ent. Soc. Amer., vol. 26, pp. 86-94, 3 figs. 
WESTWoop, J. O. 
1874 Thesaurus entomologicus Oxoniensis. 205 pp., 40 pls. 
WHEELER, GEORGE C. 
1943 The larvae of the army ants. Ann. Ent. Soc. Amer., vol. 36, pp. 
319-332, 2 pls., 2 figs. 
1948 The larvae of the fungus-growing ants. Amer. Midl. Nat., vol. 40, pp. 
664-689, 3 pls., 3 figs. 
WHEELER, WILLIAM Morton 
1919 The parasitic Aculeata, a study in evolution. Proc. Amer. Philos. 
Soce., vol. 58, pp. 1-40. 


U. S. GOVERNMENT PRINTING OFFICE: 1953 


PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1953 No. 3314 


PHOTURIS BETHANIENSIS, A NEW LAMPYRID FIREFLY 


By Frank A. McDermott 


In August 1949, at Bethany Beach in southern Delaware, some ob- 
servations were made on the few lampyrids then still flying. A single 
specimen of Photinus pyralis (Linnaeus, 1767) and one of Photuris 
versicolor (Fabricius, 1798) were observed, and a very few instances 
of a double greenish flash were noted over grass in a large vacant area 
at the north end of the town and perhaps 1,000 feet from the ocean 
front. The insects giving this flash were not captured, however, in 
spite of much pursuit, but another insect was seen to fly into the grass 
while giving the frequently repeated, rather dim flash characteristic of 
females of Photuris. This one was taken and proved to be a rather 
smal] female Photuris with a distribution of the black pigmentation on 
the pronotum different from any previously seen. Previous visits 
to Bethany Beach earlier in the year had yielded nothing notable, ex- 
cept the relative scarcity of lampyrids. In July 1951 I again spent a 
few days at Bethany Beach, and again saw the double greenish flash. 
This was considerably later than the usual period of prevalence of P. 
versicolor, and it seemed probable that this flash was that of a different 
species. Several males were taken, and one female, the latter unfor- 
tunately lost by accident, but it may be noted that the females are 
apparently relatively scarce. 

It differs from other photurids so far found in Delaware by the 
relatively small size, the distribution of the pronotal pigmentation, 
and the rather distinctly different flash of the male. The pronotal 
black pigmentation, while rather variable, has a distinct tendency to 
be enlarged toward the apical margin, and in none of the specimens 
did this spot show the forms characteristic of those of P. versicolor, 


22842353 35 


36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


P. lucricrescens Barber, 1951, or P. hebes Barber, 1951, all of which 
had been collected during the same summer in northern Delaware. 
The characteristic male flash is much like that of Photinus consangui- 
neus LeConte, 1851, two short, bright coruscations separated by a 
distinct interval perhaps somewhat longer than that of a single corus- 
cation, but is quite greenish as compared with the yellow light of 
consanguineus. Occasionally the second coruscation may be much less 
bright than the first, even with the same specimen, and a few appar- 
ently gave only a single flash but were indistinguishable from those 
giving the double flash. 

The first male specimens were taken on July 11, 1951, over the same 
field where the single female was taken in 1949, and where they were 
stili scarce, but two days later a large colony was found flying over the 
vegetation, largely bayberry, on both sides of the road leading south 
from the town, at about two miles. Trips to areas a mile or so inland 
from the beach yielded only Photinus pyralis, here very abundant at 
the time, and an occasional stray Photuris versicolor, so the new species 
seems to be a rather strictly seacoast, almost a sand-dune, form. It 
has not been seen in several visits along the southern New Jersey coast, 
particularly at Stone Harbor, but it would be somewhat surprising if 
it were not present there. 

In view of these circumstances, this insect is here described from 
the males collected in 1951, and the single female of 1949, as a new 
species. 

Photuris bethaniensis, new species 


Description.—A. rather small lampyrid for a Photuris, 9.0-10.75 
mm. long by 3.5-4.0 mm. wide, with a black pronotal mark tending 
to be densest toward the apical edge. The characteristic male flash is 
two greenish coruscations separated by a short interval. 

Heap: Width across eyes 2.1-2.45 mm. in male, 1.8 mm. in female; 
eye length 1.15-1.3 mm. in male, 0.85 mm. in female. Frons ivory to 
yellow, about 0.8-1.0 mm. wide; interocular margins usually rather 
divergent. Mandibles fairly large, 0.65-0.8 mm. across in closed posi- 
tion; proportionately larger in the female. Maxillary palpi dark 
brown, apical joint conoidal; labial palpi lighter brown, apical joint 
mitten-shaped. Clypeus dark brown, more or less tridentate. 

ANTENNAE: Dark brown, 5.0-6.25 mm. long in male, 4.25 mm. in 
female; joints may all have pale bases, or this may be limited to basal 
three joints in the female. 

Pronorum: Scutate, sometimes extended apically, 2.0-2.5 by 
2.55-3.0 mm. ; rounded posterior angles may or may not be appreciably 
produced; red pigmentation may extend to the basal edge; black spot 
of variable shape, and characteristically mainly toward the apex, or 


NEW LAMPYRID FIREFLY—McDERMOTT 37 


at least the larger portion of it apical, the longitudinal portion being 
usually merely an irregular and sometimes interrupted line, and but 
little developed along the basal margin. 

ScUTELLUM AND MESONOTAL PLATES: White or yellow. 

Eyrra: 7.0-8.0 by 1.6-2.0 mm.; ground color brown to nearly 
black; on each a white to yellow vitta, oblique, usually extending to 
about three-quarters the elytral length; wide yellowish lateral borders 
which give a slightly oval contour; narrow sutural borders. 

VENTRAL THORACIC SURFACE: Brown. 

Tereires: Tergites 1 to 6 brown; tergite 7 lighter or only partially 
infuscate ; tergite 8 pale to white. 

Srernires: 2 to 5 usually brown, sometimes becoming darker in this 
order; sternite 5 may be yellow; sternites 6 and 7 luminous in the 
male; and sternite 8 with a median posterior point, which may be very 
short. The luminous area in the female differs from the pattern seen 
in versicolor, being a transverse bilobed area medially very narrow, on 
segment 6, and a relatively large elliptical area on segment 7. 

Lres: Long; tibial spurs long, pattern 0-2-2. 

Axpracus: In the one dissected, 2.25 mm. long, identical in appear- 
ance with that of P. lucicrescens and the other photurids described by 
Barber. 

Type and paratypes —USNM 61469. 

Type locality.—Bethany Beach, Del. 

Material examined.—Holotype (male) and five paratypes, collected 
by McDermott at type locality, July 11, 1951; paratype (female), col- 
lected by McDermott at type locality, August 1949. 

Remarks.—Photuris bethaniensis is probably a seacoast form; it 
has not been noted elsewhere. The flash suggests P. fairchildi Barber, 
1951, but the latter is a larger species with narrow elytral borders. 


1 Barber, H.S. North American fireflies of the genus Photuris. Smithsonian Misc. Coll., 
vol. 117, No. 1, pp. 1-58, 3 figs., 1951. 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Voi. 103 Washington: 1953 No. 3315 


DISTRIBUTION, GENERAL BIONOMICS, AND RECOGNITION 
CHARACTERS OF TWO COCKROACHES RECENTLY ESTAB- 
LISHED IN THE UNITED STATES 


By AsHiey B. Gurney’ 


Introduction 


In contrast to the large numbers of cockroach species that occur 
in a great variety of natural habitats in most tropical countries, only 
about 55 are now recorded from the entire United States. With the 
exception of those occurring in a few localities, mainly in the South- 
western States, our roaches are well known to entomologists. In the 
northern third of the country there are few native roaches living in 
fields or woodlands, and most of the better-known species are estab- 
lished adventives which are rather closely associated with human 
habitations, having been brought there by artificial means. The recent 
establishment in the Northeastern States of two potentially important 
roaches new to this country is therefore of decided interest. 

Beginning in 1948 and continuing each subsequent year, specimens 
of E'ctobius livens (Turton), the spotted Mediterranean roach,? have 
been collected at Falmouth, Mass., under outdoor conditions, coming 
to houses at night and also occurring on fresh vegetables. Because 
of these habits and the wide natural occurrence of this roach in Europe 
and adjacent areas, it seems quite probable that a slow spread in the 
United States may take place, and that from time to time questions 
regarding its identity and importance will arise. 


1Bureau of Entomology and Plant Quarantine, Agricultural Research Administration, 
U. S. Department of Agriculture. 


2No generally accepted common name seems to be applied to H. livens in Burope, and 
this name is considered suitable for American use in view of the species’ appearance and 
the region of its dominant Old World occurrence. 
39 


228354—53——1 


40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


A second species not previously known to breed in the United States 
is Leucophaea maderae (Fabricius), the Madeira roach. Unlike the 
spotted Mediterranean roach, LZ. maderae is well known because it is 
frequently transported with articles of commerce in tropical countries. 
It has many times been intercepted by quarantine inspectors at 
United States ports and has occasionally been carried to inland cities, 
where it failed to survive. In the autum of 1950 it was found to 
be established in the basements of New York City buildings occupied 
by people who had come from Puerto Rico. Although its native 
home is evidently West Africa, it apparently was introduced to the 
West Indies more than 100 years ago. The Madeira roach could 
probably live outdoors in tropical Florida and perhaps at the other 
southern extremities of the United States, but it is more likely to be 
important as a domestic pest in the well-heated buildings of our larger 
cities. One has only to recall the increasing importance of the brown- 
banded roach Supella supellectilium (Serville) in this country during 
comparatively recent years to realize the potential importance of 
Leucophaea maderae. Apparently a native of Africa that has been 
spread by commerce to most of the warmer parts of the world, S. 
supellectilium was first reported from the United States in 1903, based 
on specimens from Key West, Fla. It was not noted as a pest here 
until found in a private home in Nebraska in 1929, but since then 
has rapidly gained recognition as one of our principal pest roaches 
(see Back, 1937, and Gould and Deay, 1940). 

This paper is intended to aid in the identification of these two 
roaches and to supply such available information on distribution and 
habits as is likely to be helpful to subsequent studies of these species 
in the United States. A third species, Vawphoeta cinerea (Olivier), 
was found to be established in Florida after this manuscript was 
prepared. It is briefly noted on page 46. 


Genus Eciobius Stephens, 1835 
Ectobius livens (Turton), 1800? (spotted Mediterranean roach) 
Fiaure 10, a—d: PuLatE 2, Ficures 3-6 


Blatta livida Fabricius, Entomologia systematicae, vol. 2, p. 10, 1793. (Pre- 
occupied by Blatta livida De Geer, Memoirs pour servir 4 l’histoire des In- 
sectes, vol. 3, p. 538, pl. 44, fig. 6, 1773.) 

Blatta lwens Turton. A general system of nature... , vol. 2, p. 529, 1800? 
(New name for Blatta livida Fabricius.) 

Ectobius lividus Fabricius, Stephens, Illustrations of British entomology, Mandi- 
bulata, vol. 6, p. 48, 1835. 

Eectobius livens (Turton), Ramme, Mitt. Zool. Mus. Berlin, vol. 27, pp. 34-45, pl. 10, 
fig. 3, 1951. 


Nomenclatural comments—Until 1951 Fetobius lividus was usually 
applied to this species. The name /. divens has been resurrected by 


COCKROACHES NEW TO UNITED STATES—GURNEY 41 


Ramme (1951, p. 34) who says that Dr. K. Princis of Lund, Sweden, 
informed him that /évida Fabricius was preoccupied and that Turton 
proposed divens on page 526, vol. 2, of his 1806 edition of Linné’s Sys- 
tema Naturae. Turton published two editions of an English transla- 
tion of the thirteenth (Gmelin) edition of this work. These editions 
are listed in the 1903-1915 Catalogue of the Books, Manuscripts, Maps 
and Drawings in the British Museum (Natural History) (vol. 3, 1910, 
p. 1128). I have verified their dates and contents by an examination 
of the entire 7-volume 1806 edition in the Library of Congress and of 
volume 2 of the 4-volume first edition in the library of the U. S. Na- 
tional Museum. The first Turton edition appeared from 1800 to 1802. 
Volume 2, containing the genus 4latia, is dated 1800 and is reasonably 
sure to have appeared no later than 1802. The second Turton edition 
appeared in 1806, the first 4 volumes being re-issues of the former 
edition with slight changes of the title pages. On the page indicated 
by Ramme (p. 526) Forficula livida, an earwig, is treated. 

Prior to Ramme’s monograph, the identities of the British species of 
Ectobius were confused. ‘Thus the species treated by Lucas (1920, p. 
78) as perspicillaris Herbst is in reality dévens. This has been clarified 
by Blair (1934, 1935). Princis (1936) has explained that a Lapland 
record of lividus (i. e., livens) actually was based on material of 
lapponicus (Linnaeus). Hebard (1948, p. 12) states that Australian 
records of lividus are incorrect. 'Taxonomists now differentiate the 
species of /’ctobius on rather minute characters, especially on the basis 
of the dorsal abdominal gland and genitalia of males. Ramme (1949) 
has described £. siculus from Sicily, differing from dévens in having 
black legs and antennae. /’. finote Chopard of Algeria (see Chopard, 
19438, p. 21) is much like livens, having the small tegminal spots between 
the veins and with the dorsal] gland bearing a rounded tubercle. The 
genus Hctobius (genotype: lapponicus) in a modern sense is much 
more restricted than formerly. In the older literature, species of 
Ectobius were reported from many parts of the world. Certain of 
these are now known to belong to other genera. For instance, many 
years ago several Nearctic roaches were referred to /'ctobius. Rehn 
(1931, pp. 806-374) and Hebard (1943) have discussed the generic 
limits of Ectobius: Rehn (p. 308) has explained the nature of the 
toothed tarsal claws; the teeth were evidently overlooked by Hebard 
(p. 12). 

Recognition features—The most important features enabling rec- 
ognition of E'ctobius livens are: (1) Small size, over-all length about 
8 to 9 millimeters; (2) small, dark spots on the veins of the front wing 
(tegmen) ; (3) conspicuous intercalated triangle at apex of wing (itr, 
fig. 10, 6); (4) claws of unequal length, the posterior claw of each 
pair much longer than anterior one; (5) in males the characteristic 
shape of the dorsal abdominal glandular depression and the sub- 


42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


genital plate (fig. 10, a, ¢) ; (6) conspicuous spines on ventroposterior 
margin of each femur. 

Of the foregoing features, the second and fifth are specific for the 
separation of this roach from other species occurring in the United 
States and the others are highly confirmatory when in combination. 

The native Nearctic species most likely to be mistaken 3 for Z. ivens 
are Chorisoneura texensis Saussure and Zehntner, of Texas and the 
Southeast, and members of the genus Cariblatta. Both have claws of 
equal length, and Cariblatta lacks a well-developed, intercalated tri- 
angle. C. texensis has a large intercalated triangle, but has no con- 
spicuous spines along the posterior margins of the femora. The best- 
known outdoor roaches of the Eastern States belong to the genus Par- 
coblatia; all are larger than divens and lack an intercalated triangle. 

Identification of the Massachusetts specimens of iévens was aided 
by comparisons made at the Academy of Natural Sciences of Phila- 
delphia with authentic French and Swiss material, given to the Acad- 
emy by the Swiss entomologist Henri de Saussure. 

Description—Avvurs: General form as in plate 2, figure 6; tegmina 
and wings covering abdomen; width between eyes on vertex about 114 
times length of first antennal segment (female), slightly less in male; 
all femora with conspicuous spines on ventroposterior margins; ven- 
troposterior margin of front femur with three or four well-spaced, 
strong spines along part slightly basad of middle, the more apical one 
often smaller than others, followed by 10 to 15 small, delicate spines 
(some of which are frequently broken) extending in a regular row 
nearly to apex, a long curved apical spine closely preceded by a straight 
subapical one about three-fourths as long; tarsal segments one to four 
each with a tiny, round pulvillus (membranous pad) ; segment 5 with 
large, broad apical arolium between conspicuously unequal tarsal 
claws, anterior claw about two-thirds length of posterior one, longer 
claw with about four spinelike inner teeth along basal half, teeth on 
smaller claw poorly developed. Tegmen with costal veins simple, 
cubital veins (discoidal sectors) strongly oblique, apex rather sharply 
rounded. Wing with distinctive intercalated triangle (itr, fig. 10, 
b). 

General color pale yellowish; tegmina and marginal area of pro- 
notum transparent, marked with reddish brown spots, on the tegmen 
these spots located on the veins; eyes dark brown to black; face yellow 
to reddish orange; vertex and upper part of face usually with several 
spots of darker orange; coxae and femora unspotted; tibiae with 
dark brown at bases of some spines; venter of abdomen with poorly 

$The standard taxonomic reference on Nearctie Blattaria is that of Hebard (1917). A 


recent key to genera is by John W. H. Rehn (1950). The European species of Hctobius 
were reviewed by Ramme (1923). 


COCKROACHES NEW TO UNITED STATES—GURNEY 43 


developed longitudinal rows of dark brown submarginal and median 
spots. 

Measurements (in millimeters) of representative specimens: Over- 
all length, including folded tegmina, female 9, male 8.5; pronotum, 
female 2.1, male 1.9; tegmen, female 7, male 7.2. Width of pronotum, 
female 3, male 2.9. No significant size variation has been noted. 

Nymrus: Nymphs (pl. 2, figs. 3, 4) ranging in body length from 
2.5 to 4.5 millimeters have been examined. They are yellow, with 
conspicuous dark reddish brown spots on the dorsal surface and on the 
vertex and upper part of the face. Dark submarginal longitudinal 
streaks appear on the thorax. The venter of the abdomen is dark 
brown submarginally and medially. 

OdruEcAE: Two odthecae have been examined (pl. 2, fig. 5). They 
are dark brown and measure 2.3 millimeters in length. The convex 
margin is weakly crenulate and there are faint transverse indications 
of the eggs contained. 

Distribution—Three American localities are known for E'ctobius 
livens, all in Massachusetts: Falmouth, Manomet, and Plymouth. 
Falmouth, the first site discovered here, is situated near Woods Hole 
and the Cape Cod Canal, on the southern side of the base of Cape Cod. 
In October 1951, after this manuscript had been prepared, I ex- 
amined 3 males and 4 females of livens collected at Manomet, Plym- 
outh County, Mass., by Mrs. S. P. Graeff and submitted by Dr. Ells- 
worth H. Wheeler of the University of Massachusetts. Manomet is 
on the coast of Cape Cod, about 20 miles north of Falmouth. The 
specimens were collected before August 13, 1951, apparently just 
prior to that date. Mrs. Graeff wrote, “Our summer cottage is pretty 
well overrun with them. Painters told me they were even on the roof. 
The cottage is in the woods and the underbrush is close. I think this 
is the third summer we have had them. I suspect they like damp 
spots, they scurry under shingles.” 

An adult of livens was submitted July 28, 1952, by a Plymouth 
housewife who reported that her house and shrubbery were becoming 
infested. 

This roach is widely distributed in the southern and central portions 
of western Europe, occurring in Germany, the Netherlands, Belgium, 
France, Switzerland, Italy, Portugal, Spain, and southern England. 
It has also been reported as far eastward in the Mediterranean area 
as Asia Minor, but the records I have seen were published prior to 
the modern work dating from Ramme’s 1923 monograph of Letobzus 
and are not dependable. Chopard (1948, p. 21) has reported it from 
Algeria and Tunisia. I have found no records of its occurrence in 
the Azores. Fabricius originally described the species from France, 
and, in the absence of type material, Ramme in 1923 designated a male 


44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


neotype from Montpellier in southern France. Two varieties of 
livens have been recognized, chopardi Adelung described from France 
in 1916, and mznor Ramme described from the Island of Elba in 1923. 
These varieties are based primarily on the amount of dark color on 
the lower surface of the abdomen, minor being quite dark except near 
the lateral margins, only the submarginal dark spots being well de- 
veloped in chopardi, and the submarginal spots being poorly or moder- 
ately developed in typical Zivens. The species of Hctobius are well 
known for color variation, the color of tegmina and abdomen often 
being recessive (pale) or intensive (dark). In view of this situation, 
it remains to be proved whether chopardi and minor are valid sub- 
species or if they are forms without real significance. The center of 
distribution for the genus /’ctobius, as outlined by Rehn (1931, pp. 
314-318), was apparently the Mediterranean Basin, with a secondary 
center developed in Africa south of the Sahara Desert. 

Biology.—The original Massachusetts specimens of Hctobius livens 
examined consist of 2 males, 16 females, 9 nymphs, and 2 odthecae. 
The males were taken June 21, 1948, and the other specimens in late 
August and September 1950.4 It may be significant that males were 
found only in the spring and early summer. Lucas (1928) reported 
that in England males of /. lapponicus disappeared about midsum- 
mer, the females persisting longer than males, and nymphs appearing 
in late summer and fall preparatory to hibernation. No detailed 
life history studies of livens in Europe have come to my attention. 
Brown (1952) has reported observations on the related 2’. panzeri 
Stephens in England, which occurs there mainly on sand dunes near 
the coast. Adults die in the fall, the males disappearing first, but 
nymphs overwinter and do not reach full maturity until early August 
of the following year. 

The Falmouth collections were made on a farm located beside a 
small salt river about a mile from the shore. Mr. Flint found speci- 
mens in a large variety of situations, specimens occurring most con- 
sistently beneath loose lichens on oak trees, and crawling on trees and 
houses at night—the latter in the vicinity of lights. Some were 
found under baskets, in buckets, or on Swiss chard and other vege- 
tables, or were taken by general sweeping (see Flint, 1951). 

In England £. dévens occurs only in the extreme southern counties— 
there found on trees, among bracken ferns, under dead leaves, and 
flying actively in hot sunshine (Lucas, 1920, p. 80; Burr, 1936, p. 44). 
Chopard (1947, p. 37) says dévens is very common in the French woods, 
and he gives a colored illustration (pl. 2, fig. 28). 

4 All of this material was collected by Oliver S. Flint, Jr., a student at the University 
of Massachusetts. I am much indebted to Mr. Flint for the enthusiasm with which he 


sought additional material once the character of his initial captures was pointed out to 
him, as well as for his diligence in seeking clues to the origin of the introduction. 


COCKROACHES NEW TO UNITED STATES—GURNEY 45 


Source of introduction—The Massachusetts colony of H'ctobius 
livens is probably the result of an accidental introduction from western 
Europe or the Mediterranean area. A good many people of Portu- 
guese ancestry live in Falmouth and the vicinity, and visits to Portu- 
gal and the Azores are frequent. Seeds and occasional shipments 
of fruit are brought back, but quarantine regulations prevent wide- 
spread movements of plant products. Since livens occurs in Portugal, 
as reported by Seabra (1942, p. 19), the possibility of an introduc- 
tion from that part of the Mediterranean seems the most likely. 

Probable importance.—Ectobius livens is not likely to develop into 
a major pest or to live in buildings except occasionally. As a nuis- 
ance pest in the vicinity of gardens and dwellings, it may be expected 
to attract attention at intervals and to be submitted to entomologists 
for identification and advice, as is now true of male specimens of 
Parcoblatta which are attracted to lights and which occasionally re- 
main in houses for short periods. 

So far as known, species of Hctobius are not economically impor- 
tant in Europe. /. lapponicus has often been stated to be a pest of 
dried fish in Swedish Lappland, but Gaunitz (1935, 1986) found 
that lapponicus does not occur indoors and probably feeds on 
vegetable matter. 


Genus Leucophaea Brunner, 1865 
Leucophaea maderae (Fabricius), 1793 (Madeira roach) 
FIcureE 10, e-h ; PLATE 2, Fiecurss 1, 2 


Blatta maderae Fabricius, Entomologia systematicae, vol. 2, p. 6, 1793. 
Leucophaea maderae (Fabricius), Rehn, Trans. Amer. Ent. Soc., vol. 29, p. 283, 
1908. 

Nomenclatural comments.—The name Rhyparobia maderae is cur- 
rently used by certain workers, especially Europeans, as exemplified 
by Chopard (1948, p. 45) in his monograph of North African Orthop- 
tera. Leucophaea was proposed by Brunner (1865, pp. 272, 278) 
as a subgenus of Panchlora, with four included species. One of the 
four, maderae, was designated type of Leucophaea by Rehn (1903, 
p. 282). Rhyparobia was proposed by Krauss (1892, p. 165), mono- 
typic for maderae. Thus Rhyparobia is an isogenotypic synonym of 
Leucophaea. The use of Rhyparobia was doubtless encouraged by the 
action of Kirby (1904, pp. 150-151) who, evidently unaware of Rehn’s 
1903 designation, designated Blatta surinamensis Linnaeus type of 
Leucophaea. As explained by Hebard (1917, p. 309), Kirby also ap- 
parently objected to the name Pycnoscelus Scudder, 1862, to which 
surinamensis belongs, because it was originally based on an immature 
specimen. Kirby’s latter view is not supported by the rules of nomen- 


46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


clature but, regardless of that aspect of the matter, the validity of 
Leucophaea, with maderae its type, is clear. 

Recognition features—The most important features in the recog- 
nition of Leucophaea maderae are: (1) Large size, overall length about 
38 to 53 millimeters; (2) pronotal shape, and general color markings 
as illustrated (pl. 2, figs. 1, 2); (8) no strong spines along posterior 
margins of femora except one short apical spine on each hind and 
middle femur; (4) in males the characteristic shape of the specialized 
organ on the dorsal surface of the second abdominal segment and of 
the subgenital plate (fig. 10, e, A.). 

Relatively few Nearctic roaches are as large as L. maderae, and none 
of them is likely to be confused with it. Blaberus craniifer Burmeister, 
illustrated by Hebard (1917, pl. 8, fig. 6), is the largest roach in the 
United States, being much larger than maderae. The species of 
Periplaneta are quite differently marked, and have numerous heavy 
spines on the posterior margins of the femora. Both Furycotis 
floridana (Walker), of the Southeast, and Hemiblabera tenebricosa 
Rehn and Hebard,’ of the Florida Keys, have short, lobate tegmina 
and wings which are vestigial or lacking. 

A relative of Z. maderae recently established in Florida is 
Nauphoeta cinerea (Olivier). It superficially resembles maderae, 
though it is smaller, its average length being about 23 millimeters. 
The femora are proportionately stouter and the tegmina scarcely ex- 
tend beyond the apex of the abdomen, while the posterior margin of 
the pronotum is transversely subtruncate, a median projection being 
scarcely evident. Zimmerman (1948, p. 94) and Rehn (1945, p. 274) 
have given photographs of cinerea. This species is apparently a na- 
tive of East Africa now widely distributed in the Tropics by com- 
merce and often intercepted at United States ports. Its potential 
importance is suggested by Illingworth (1942), who described its oc- 
currence in alarming numbers about the feed room of the poultry 
plant at the University of Hawaii. He also mentioned its occurrence 
in heated buildings in England and Germany, though no records of 
permanent establishment in those countries have come to my attention. 

In early December 1951 material of cinerea was received from Jo- 
seph Gross of Tampa, Fla., who stated that it was widely and thor- 
oughly established about Tampa, especially in feed mills. Later, in 
August 1952, Mr. Gross contributed a series of 34 adults and 32 nymphs 

5 Hemiblabera tenebricosa was described from the West Indies, including Nassau in the 
Bahamas, by Rehn and Hebard (1927, p. 271). Caudell (1931) recorded an adult pair 
collected at Key Largo, Fla., by E. A. Popenoe in 1896. Caudell’s unpublished notes dis- 
close subsequent discussions of this record with Mr. Popenoe’s son, who explained that his 
father was collecting insects primarily for Kansas State College during the 1896 trip and 
that in view of conditions at Key Largo at that time this roach was probably an established 
species rather than an intercepted adventive. A number of roaches are now known to 


be native to the northern Bahamas and extreme tropical Florida, so the occurrence of 
tenebricosa in the Florida Keys 1s not illogical. 


COCKROACHES NEW TO UNITED STATES—GURNEY 47 


Ficure 10.—Structural details of Ectobius livens (Turton) and Leucophaea maderae (Fab- 
ricius). a-—d, E. livens: a, Dorsal view of apical half of cleared male abdomen; b, wing; 
c, ventral view of subgenital plate and associated cerci, male; d, apical portion of genital 
hook, male. e-h, L. maderae: e, Dorsal view of second abdominal segment, male, cleared 
and in alcohol; f, dorsal view of apical half of cleared male abdomen; g, genital hook of 
male, enlarged; f, ventral view of subgenital plate and associated structures, male, dry 
preparation. Drawn from specimens as follows: a—d, from Massachusetts specimen; e, 
from Puerto Rico specimen; f and g, from specimen collected in a Tampa, Fla., warehouse; 
h, from New York City specimen. (A-9, apodemes of sternum 9; ca, oblique carinae of 
glandular specialization; gh, genital hook; gld, glandular depression; itr, intercalated 
triangle; me, mesal extension; st, stylus; sap, supra-anal plate; sgp, subgenital plate.) 

228354—53 2 


48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


to the U.S. National Museum. I inquired especially as to the apparent 
permanence of the infestation, and on June 24, 1952, a colleague of Mr. 
Gross’, W. B. Gresham, Jr., replied, “It seems evident to us that 
Nauphoeta is in Florida to stay. They appear to be well adapted and 
quite numerous in the locations where we have noted them. All in- 
festations noted have been within commercial buildings engaged in the 
manufacture of animal feeds. I myself have not noted any out of 
doors.” Notes on the Florida infestations by Ratcliffe (1952) and 
Gresham (1952) have appeared. 

Description.—Avutts: General form as in plate 2, figure 1; tegmina 
and wings extending beyond apex of abdomen about one-fifth their 
length; width between eyes on vertex about two-thirds length of first 
antennal segment; a shallow, scarcely wrinkled transverse depression 
in interocular area where eyes are closest; ventroposterior margin of 
middle and hind femur each with short, apical spine; conspicuous 
spines on posterior femoral margins otherwise lacking; ventropos- 
terior margin of front femur with continuous row of delicate, seta- 
like spines, these slightly shorter toward apex of femur and a few 
noticeably longer ones near base; front tibia with two elongate groups 
of setae along margins; pulvillus on each tarsal segment, elongate 
on segments 1 and 5; claws equal, unarmed ventrally, with sparse 
setae dorsally ; pronotum obtusely angulate laterally, posterior margin 
with rounded median production; tegmen with cubital veins (dis- 
coidal sectors) oblique; wing (see J. W. H. Rehn, 1951, pl. 9, fig. 102) 
with well-developed axillary and cubital fields, intercalated triangle 
lacking. Dorsal surface of abdomen simple in female, specialized in 
male; second tergum of male (fig. 10, ¢) with median glandular de- 
velopment consisting of paired oblique carinae (ca) and a brief longi- 
tudinal carina centered on a trifid papilla which largely covers a 
small glandular depression (gld) ; supra-anal plate transverse, with 
median emargination of posterior margin, general shape more quadrate 
when dry (fig. 10, 2) than when preserved in alcohol (fig. 10, f) ; sub- 
genital plate unspecialized in female, slightly asymmetrical in male, 
with recurved lateral hooks near styli; cleared male abdomen with 
apodemes of sternum 9 (subgenital plate) (A9) united anteriorly, 
genital hook (gh) and additional sclerotized elongate rod associated 
with an irregular genital mass; genital hook (fig. 10, g) enclosed in a 
membranous sheath. 

General color pale brown, the tegmen and pronotum marked with 
dark brown as illustrated (pl. 2, fig. 1) ; remainder of body and ap- 
pendages largely dark brown; clypeus and most of labrum pale; ab- 
dominal sterna with oblique submarginal marks of brownish-black; 
terga with more heavily pigmented areas as shown (fig. 10, 7). Base 
of genital hook with pigmented transverse lines, the apical portion 
much darker. 


U, S, NATIONAL MUSEUM PROCEEDINGS, VOL. 103 PLATE 2 


1,2, Leucophaea maderae (Fabricius); 1, Adult female, collected in Manila, Philippine 
Islands (length, 47 mm.): 2, last nymphal stage, intercepted at Washington, D.C. 

3-6, Ectobius livens (Turton), all from Falmouth, Massachusetts: 3, 4, Nymphal stages; 
5, odtheca; 6, adult female (length, 9 mm.). 


COCKROACHES NEW TO UNITED STATES—GURNEY 49 


Measurements (in millimeters): Overall body length, including 
folded tegmina, female 45 to 53, male 38 to 45; pronotum, female 9 to 
10.3, male 7.5 to 8.5; tegmen, female 37 to 46, male 32 to 38.5. Width 
of pronotum, female 13.3 to 16, male 11.5 to 14. 

Nympus: Nymphs (pl. 2, fig. 2) ranging from 7.5 to 82 millimeters 
in body length have been examined. The dorsal surface of the thorax 
and abdomen is sharply rugose, with short, microscopic spines, these 
especially conspicuous along the posterior margins of segments; 
much smaller and less evident spines on ventral surface of abdomen. 
General color reddish brown; each tergum with a submarginal dark 
spot near the base, the basal half of the exposed lateral margin black- 
ish and the apical third yellowish. 

Distribution—In the United States Leucophaea maderae is known 
to be established only in the Harlem section of New York City.° Many 
times each year inspections made at shipping centers detect the species 
with plant products and other articles coming both from the American 
Tropics and the Old World. The distribution of maderae in warm 
countries is very wide, but somewhat irregular, depending upon the 
vagaries of commerce. The specific and the common names are de- 
rived from the fact that this roach was first described from Madeira, 
a small island now belonging to Portugal and located in the Atlantic 
Ocean about 400 miles west of Morocco. Rehn (1937, pp. 56-58; 1945, 
p. 273) has reasoned (by a detailed analysis of the present occurrence 
of maderae and its congeners) that West Africa is the native home, 
from which commerce (in early centuries—that of the slave trade in 
particular) has carried it to Madeira, the West Indies, Brazil, and 
elsewhere. Ecuador, Colombia, Venezuela, the Guianas, Brazil, and 
Argentina are known to have established colonies of maderae, as well 
as Panama, Costa Rica, and most of the West Indian islands. Old 
World distribution outside of Africa is more spotty and, to my knowl- 
edge, the species is not yet recorded from India or Australia—even 
though it is known from Java, the Philippines, Hawaii, and Fiji. 
Doubtless, maderae now breeds in many coastal areas from which 
there are no records and it will continue to spread as conditions permit. 

Biology.—In New York City Leucophaea maderae has been “found 
in some abundance in the basement of an apartment of the Harlem 
section,” and an observer also writes? “From their abundance and the 
presence of all stages of the insect . . . . I would conclude that they 


®Davis (1940) reported at a meeting held December 15, 1938, that a specimen of 
ZL. maderae taken in the Bird House of Bronx Park, New York City, was the first record 
of this species in New York State. In the absence of further information, there is a strong 
likelihood that the specimen in question was an escaped adventive. 

7 Specimens from the described infestation were submitted by Ralph E. Heal, Technical 
Director of the National Pest Control Association. Grateful acknowledgment is made of 
Dr. Heal’s cooperation in obtaining information on the Harlem infestation. 


50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


are breeding in this house.” It was also noted that the Harlem roaches 
appeared rather sluggish in their movements. 

In the absence of biological studies of Z. maderae in this country it 
is instructive to review the observations of life history and habits 
that have been made elsewhere, particularly the papers of Illingworth 
(1915), Sein (1923), and Pessoa and Correa (1928). 

As is true of species of Periplaneta, growth in the Madeira roach 
isslow. The first molt may occur 1 to 4 months after birth. Pinned 
nymphs reared by Illingworth in Hawaii and now deposited in the 
U. S. National Museum suggest that there are six nymphal stages, 
though the number is likely to vary, and in cultures it may sometimes 
be difficult to observe because exuviae reportedly are eaten immedi- 
ately after each molt. Pessoa and Correa report only four “meta- 
morphoses,” but that number of molts is almost certainly too small 
for a normal growth cycle. The body length in millimeters of speci- 
mens of various ages collected by Illingworth is as follows: 1 day, 
7.5; 1 month, 7.5; 4 months, 16; 5 months, 18.5; 6 months, 24.5; 7 
months, 29; 8 months, 30.7; 9 months, 32. Food and environmental 
conditions are doubtless important influences on growth. Sein notes 
maturity being attained in 220 days, but that a year is often required, 
while Pessoa and Correa indicate that adults are obtained in a year. 
with 16 to 18 months the longest observed time. In Sein’s cultures 
the first nymphs appeared 4 months after their parents matured. 
Immediately following birth, nymphs usually hide beneath the 
mother during the day. According to Pessoa and Correa, this makes 
the adult restless and active in contrast to its usual slow gait. 

In a general paper emphasizing the adaptability and value of 
Leucophaea maderae as an experimental animal, Scharrer (1951) 
comments that it thrives on a diet of apples, carrots, and dogfood. 
She reports that there are 30 to 35 young every 8 months, that there 
is an average of 8 molts, and that life expectancy is up to 214 years. 
Dr. Scharrer has observed that tumors which resemble malignant 
cancer of higher animals develop in various parts of the roach body 
following the removal of the corpora allata and the corpora cardiaca. 
That such an injury to the nervous system has produced tumors has 
very interesting and practical implications for further research. 
The name “woodroach” that Dr. Scharrer applied to maderae is a 
questionable choice, since the name is most often used for native 
Nearctic species of Parcoblatta. 

Leucophaea is one of several roach genera which are viviparous; 
that is, the eggs are enclosed in a delicate membranous sac which nor- 
mally ruptures before extrusion from the mother’s body, or immedi- 
ately thereafter, so that, in effect, the young are born alive. This is 
in contrast to the habit of most roaches which form a heavily-sclero- 
tized, dark-colored capsule or odtheca, usually of characteristic shape, 


COCKROACHES NEW TO UNITED STATES—GURNEY 51 


which may be carried protruding from the body for some time prior 
to hatching, or be deposited loosely or surrounded with a matrix. 

The viviparity exhibited generally takes a special form known as 
ovoviviparity. This occurs in several well-known genera, especially 
Panchlora, Pycnoscelus, Nauphoeta, and Leucophaea, and Chopard 
(1938, p. 218) notes six subfamilies of roaches (according to the long- 
used classification) in which viviparity occurs. J.W.H. Rehn (1951)' 
has recently proposed a new classification, based on wings, and it is 
significant that many of the viviparous genera fall in what he terms 
the epilamproid complex. In addition to Chopard, Shelford (1907), 
Karny (1924, pp. 3-10), and Rau (1941) have discussed viviparity 
among roaches. Hagan (1941; 1951) has described the female re- 
productive system of the viviparous Diploptera, and Chopard (1950), 
that of Gromphadorhina. 

Illingworth and Sein both recognized that Leucophaea maderae is 
viviparous, but it appears that Pessoa and Correa observed unusual 
or abnormal instances of the egg sac being deposited, as they wrote of 
a capsule being placed in the darkest corner of a rearing box and the 
first young appearing 20 days later. In the following paragraphs I 
have attempted to explain this apparent lack of agreement. 

Several preserved adult females of Z. maderae received for identifi- 
cation during recent years have exhibited an elongate sac about 20 
millimeters long protruding from the end of the abdomen. In cer- 
tain cases the eggs (varying up to about 40 in number), directed 
transversely with respect to the mother’s body and arranged in two 
rows, were undeveloped, but in others nymphs ready for hatching were 
visible—in fact, rupturing of the sac had sometimes occurred. Rehn 
(1937, p. 62) has described his experience in the Belgian Congo with 
the related Z. grandis (Saussure). A female confined in a bottle gave 
birth to 20 living young. The number of young of maderae produced 
at one time, according to published records, ranges from 25 to 32. 

The related Pycnoscelus surinamensis (Linnaeus) is known to be 
viviparous, but the literature concerning its habits suggests a lack of 
uniformity in birth or hatching. Watson (1929, p. 58) commented on 
an egg capsule being deposited but the eggs failing to hatch. Caudell 
(1925) also maintained cultures of P. surinamensis, and mentioned 
young born alive and odthecae seldom if ever protruding from the 
female abdomen. In a somewhat fuller account, Zappe (1918) gave 
the opinion that young of swrinamensis are either born alive or hatch 
from eggs within 24 hours. Eggs were often laid in soil, but were not 
observed to hatch. When females were injured or excited, they often 
deposited poorly developed egg masses that did not hatch. Similarly, 
Illingworth (1942) reported that the eggs of Nawphoeta cinerea were 
usually kept in the body until living young appeared, but that confined 
individuals sometimes produced aborted egg masses that were ex- 


52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


truded but which dried up without hatching when the delicate enclos- 
ing membrane was exposed to the air. These experiences suggest that 
some of the protruding eggs of Z. maderae that I have noted are the 
result of handling when the specimens were collected and that occa- 
sional egg masses may be deposited under certain conditions by this 
ordinarily viviparous species. 

In his important recent paper, Chopard (1950) has discussed the 
anatomy and development of Gromphadorhina laevigata Saussure 
and Zehntner, of Madagascar. He observed, in the course of rearings 
conducted in Paris, that the female of this viviparous species extrudes 
the soft odtheca nearly to its full extent, then it is drawn into an 
incubating pouch where the eggs undergo incubation for approxi- 
mately 70 days prior to the appearance of the young. Occasionally 
the odtheca is completely extruded, with the result that desiccation 
occurs and no hatching takes place. I am indebted to my colleague 
R. E. Snodgrass for pointing out the significance of Chopard’s obser- 
vations. They explain how the oétheca is transferred from the uterus 
to the incubating chamber in the case of Gromphadorhina. Possibly 
the same habit occurs in certain other viviparous genera, and some 
protruding odthecae that have been seen may represent a stage in this 
normal act. In discussing the transfer of the developing eggs in 
the viviparous Diploptera, Hagan (1951, p. 299) states that the 
odcytes pass from the ovarioles to the lower end of the common ovi- 
duct, where “they are directed by the ovipositor from the genital 
chamber ventrally into the open end of the uterus.” 

In Brazil, mating of Leucophaea maderae occurs mainly during the 
warm and rainy season, according to Pessoa and Correa, who say that 
copulation may occupy 20 to 30 minutes and takes place with the pair 
end to end facing in opposite directions. Prior to mating, the female 
is described as opening her wings and drawing them along the ground, 
at the same time producing a sound by vibrating them. The scent 
gland on the dorsum of the male presumably is attractive to the female 
at mating time, but its function is not definitely known. 

Illingworth stated that when disturbed the Madeira roach stridu- 
lates very noticeably, and he believed the sound is produced by rubbing 
the posterior margin of the pronotum over the mesonotum. Accord- 
ing to Chopard (1938, p. 286), several roaches stridulate delicately by 
rubbing the border of the pronotum upon the mesonotum or upon the 
strongly denticulate base of the costal vein of the tegmen. To test the 
possibilities of stridulation, I have relaxed dry specimens of both sexes 
of L. maderae, then manipulated the body parts with my fingers. A 
low, squeaking sound is consistently obtained in either sex by rubbing 
the lateroposterior margin of the pronotum on the basal costal margin 
of the tegmen. Both surfaces are heavily sclerotized, turned to oppose 


COCKROACHES NEW TO UNITED STATES—GURNEY 53 


each other, nonpubescent, and are finely rugose so as to make stridula- 
tion possible. 

In tropical regions where it is established, ZL. maderae is definitely a 
domiciliary species, though, like certain other roaches often associated 
with man, it is capable of living apart from him in a purely wild state. 
From available records, however, it is not clear how frequently it has 
been collected entirely unassociated with man-made surroundings. 
According to Alfken (1904, p. 565) more than 50 years ago a German 
collector, H. Schauinsland, in 1896 or 1897 found maderae in native 
huts on the Hawaiian Island of Molokai, commerce already having 
carried it to certain Pacific areas. However, in spite of this long estab- 
lishment in the Hawaiian group, no mention of its occurrence in fields 
is made by Williams (1931). In July 1950, in the Venezuelan State of 
Aragua, Dr. Ernst Schwarz ® collected it coming to lights in fair num- 
bers, but whether it was breeding in the adjacent forest or associated 
with nearby habitations is not known. Likewise, specimens collected 
by the late C. F. Baker in the Philippines before 1927 bear the collect- 
ing label “Mt. Maquiling,” and they may have bred under entirely nat- 
ural surroundings. However, as I saw during my visit to Mt. Maquil- 
ing in 1945, at the base and on the lower slopes there are ample oppor- 
tunities for this roach to remain associated with man. The species 
frequently flies actively and has often been taken at lights, including 
those on porches, in field camps, or of automobiles. On the other hand, 
Sein states that Leucophaea maderae does not fly much when indoors 
at night as part of an infestation. JZ. puerilis Rehn, considered its 
closest relative, is a strictly endemic, forest type in West Africa. 

Source of introduction.—The New York City infestation is reported 
to be localized in apartment buildings occupied by people from Puerto 
Rico and, since there has been a good deal of movement—much of it 
by air—from that island in the past few years, there is little doubt that 
Puerto Rico is the source of the infestation. 

Probable importance.—In Puerto Rico the Madeira roach most often 
occurs in fruit stores and markets. It is especially fond of grapes. It 
is considered very gregarious and develops large, localized colonies. 
Wolcott (1950, p. 43) records about a bushel having been swept from 
one store. Warehouses and other buildings are often infested. When 
handled or otherwise disturbed, Leucophaea maderae produces an odor 
described as especially offensive. Houses infested by the species usu- 
ally do not contain Periplaneta or Blattella. There is a strong possi- 
bility that maderae, if allowed to spread, will gradually develop into a 
serious pest in our larger cities. In the light of its wide occurrence in 
the Bahamas and other West Indian islands, tropical Florida would 
probably be a suitable habitat, either outdoors or in buildings without 
central heating. 


8 Associated with the Venezuela Plague Mission (Commander J. M. Amberson, U.S.N., 
Dr. Ernst Schwarz, and Mrs. Schwarz). 


54 PROCEEDINGS OF THE NATIONAL MUSEUM VOI. 108 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 


U. S. NATIONAL MUSEUM 


Vol. 103 Washington : 1953 No. 3316 


= 


BITING MIDGES OF THE HELEID GENUS STILOBEZZIA IN 
NORTH AMERICA 


By Wiis W. Wirts' 


Stilobezzia Kieffer, 1911, is a rather large, widespread genus of 
predaceous midges which has reached its greatest development in 
the warmer parts of the New World. For North America 11 species 
are listed in Johannsen’s recent list (1943b); this total is now raised 
to 18 by the present new records and species, and no doubt many 
more await discovery. The species of Stilobezzia are rather easily 
characterized, and all the Nearctic species known to Johannsen were 
correctly placed by him, with the exception of S. wncinata Johannsen, 
1943, which I believe to represent the female of Parabezzia petiolata 
Malloch, 1915. 

The types of the species described here are in the U. S. National 
Museum, the collections of which furnished the bulk of the material 
studied. I am indebted to Dr. Henry K. Townes, of North Carolina 
State College, for the loan of specimens from his collection, and to 
Mrs. Elisabeth C. Beck and J. A. Mulrennan, of the Florida State 
Department of Health, for their kindness in sending a large series of 
light trap collections from which I sorted most of the Florida specimens. 


Family HELEIDAE 


Genus Stilobezzia Kieffer, 1911 


Diagnosis——Body rather slender and nearly bare. Eyes bare. 
Female antenna with segments 3-10 oval, 11-15 long and cylindrical; 


1 Bureau of Entomology and Plant Quarantine, Agricultural Research Administration, U. 8. Depart* 
ment of Agriculture. 
232991—53——_1 57 


58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


male antenna with well-developed plumes. Mesonotum usually with 
few or no long bristles; humeral pits present but usually inconspicuous. 
All legs slender; femora without spines or (in Hukraiohelea) with a 
few on forelegs; basitarsus and fifth segment with or without ventral 
spines; fourth segment cordiform, fifth not enlarged; female claws 
large and very unequal or a single long claw with basal barb on each 
leg, male claws small and equal; empodium absent. Wing rather 
long; microtrichiae present; macrotrichiae present toward tip (in 
Neostilobezzia) or absent; costa extending to two-thirds wing length 
or more; usually two radial cells, second much larger than first, 
which may be entirely absent (in Hukraiohelea) or greatly reduced; 
intercalary fork distinct; crossvein r-m more or less oblique; median 
fork with long petiole, M2 not interrupted at base; anal vein straight, 
not interrupted in middle. Male genitalia with basistyles simple or 
with lobe on inner margins; dististyles long; ninth sternite very short; 
ninth tergite usually rounded caudad with setigerous apicolateral 
lobes; aedeagus reduced to a pair of oblique lateral sclerites; parameres 
a pair of large, highly sclerotized, greatly modified, submedian proc- 
esses with flaring basal apodemes. 


Keys to the Nearctic species of Stilobezzia 
Females 


1. Only one long radial cell present; fore femur with 2 or 3 ventral spines 


(subgenus Hukratohelea)_..___.._------ 1. elegantula (Johannsen) 

Two radial cells present; fore femur unarmed_____.-__---_-----_-- 2 

2 (1). Wing with macrotrichiae at tip (subgenus Neostilobezzia)__________- 3 
Wing entirely bare (subgenus Stilobezzia)_..._._.._-...--_=---_-- 5 

3 (2). Body hairs very strong and erect; wings hyaline; legs, scutellum, and 
halteres-yelloweo8) 2325 ok ee 3. lutea (Malloch) 

Body hairs soft and decumbent; midlegs and hind legs brown_-_-_--_-_- 4 


_ 4 (3). Wing infuscated brown; scutellum and halteres yellowish. 
4, stonei, new species 


Wing hyaline; scutellum and halteres brownish... 2. fuseula Wirth 
5 (2). Wing with 2 or 3 dark spots on anterior margin______________2___1- 6 
Wing hyaline;-unmmarked.____.2_.. -..- 32522222...) eee 9 


6 (5). Shining yellow; mesonotum polished yellow, with pair of longitudinal 
brown bands, anterior spine absent; legs yellow, narrow apices of hind 
femur and tibia brown; wing hyaline, with brown spots at crossvein 
and apex of second radial cell___._____-._____ 9. eoquilletti Kieffer 

. Mesonotum brown with silvery, pruinose areas and prominent anterior 
spine; legs extensively brown banded or punctate; wing brownish, 
anterior margin with 2 or 3 dark spots /_-_=__-__/_. | 32 ee See | 

7 (6). Femora and tibiae extensively yellow, with small brown spots at bases of 
hairs; wing hyaline, with two brown spots; abdomen mostly yellowish 
BDOVE! 25 eco se ee ee 14. punctipes, new species 

Femora and tibiae brown with yellowish rings; wing infuscated, with 2 
or 3 brown spots and a subapical pale area; abdomen black above with 
narrow, distal, silvery pruinose, segmental bands__._-_.--------- 8 


S.,.(7) 


HO... (9):. 


11 (10). 


12 (10). 


13: (9). 


14 (13). 


15 (13). 


16 (15). 


17 (16). 


Z(t). 


3 (Ll). 


4 (3). 


HELEID MIDGES, GENUS STILOBEZZIA—-WIRTH 59 


Three dark wing spots including one at apex of first radial cell; fifth tarsal 


segment with ventral batonnets__-_------------ 15. rabelloi Lane 
Two dark wing spots, none at apex of first radial cell; fifth tarsal segment 
without ventral batonnets__-.------------ 6. beckae, new species 

we Vesonotumuentirely polishediblack 22-2 22 === 4-5 5ss5 5-522 552 10 
Mesonotum yellow, green, or brown, not entirely black____-_----- 13 
Abd omen polishediblack=s 25-5 = 545 25252 8 ae eee ee 11 
PINCH CRM OTERO 25-5 2ta Ee eR hee ee ae es it 12 


Midfemora and hind femora black except at extreme bases. 

5. antennalis (Coquillett) 
ihepsuinifermiy, yellow@4.10 829 5-bc eke eee 2 7. bicolor Lane 
Fifth tarsal segment with ventral batonnets__10. diversa (Coquillett) 
Fifth tarsal segment without ventral batonnets. 

12. pallidiventris (Malloch) 
Shining green; third or fourth abdominal tergites extensively black. 14 
Yellow or brown, not green; abdomen without black bands or spots. 15 
Mesonotum uniformly pale green; distal fourth of hind femur black. 
18. viridis (Coquillett) 
Mesonotum largely dark brown; hind femur with only spot at apex 
| OVENS} ice Re 2 oe Ne a me Oe FEL Be Ore ee ee 11. glauca Macfie 
Mesonotum shining brown; wing with a beadlike swelling near apex of 
second, radial cell 22 2G eek Be Se 16. sybleae, new species 
Mesonotum pruinose brown; wing venation normal_-_-_-_---------- 16 
Mesonotum dark brown, with pattern of large pruinose gray spots; 
scutellum dark; tibiae with sub-basal pale rings_-13. pruinosa Wirth 


Mesonotum uniform light grayish brown; tibiae unbanded_-----_-- 17 
Scutellum and femora yellow___-.---------------- 8. bulla Thomsen 
Scutellum and femora brown---------- 17. thomsenae, new species 


Male genitalia 


Lateral sclerites of aedeagus greatly reduced or absent, instead a trans- 


verse sclerite between inner margins of basistyles beyond base_ - - _- 2 
Lateral sclerites of aedeagus well developed and oblique, with bases at 
TN CE VS UGTA 01 OTS Oly LD EVSIS ty OS ee ee eee 3 


A large rounded lobe at distal third of inner margins of basistyles; 
transverse sclerite with pointed, curved ends; apicolateral lobes of 
ninth tergite widely spaced and triangular_----_._ 8. bulla Thomsen 

Basistyles simple; transverse sclerite with blunt ends; apicolateral lobes 
fingerlike and close together_______- 17. thomsenae, new species 

Lateral sclerites of aedeagus sinuate, irregularly bent, or apex forked; 
basistyle without lobe, or if one present it bears a strong spine; apico- 


lateral lobes of ninth tergite usually slender and close together _ -___ 4 
Lateral sclerites of aedeagus straight, at least on distal half; basistyle 
with mesal lobe; apicolateral lobes usually low and rounded - - --_- 10 


Basistyle with a strong spine midway on inner margin, arising from a low 
lobe also bearing a fine hair; aedeagus with strong subapical tooth 
on outer side, appearing forked at end; parameres long and slender 
EEE ONC DOLIULCOMELDS l= = = toe ee ene ee cam 2. fuscula Wirth 

Basistyle simple; aedeagal sclerites without strong tooth__. ._-_----- 5 


60 


10 


12 


16 


(4). 


(5). 


(6). 


COE 


(6). 


(3). 


(10). 


(11). 


(11). 


(13). 


(13). 


(15). 


PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Parameres greatly expanded from bases, with obliquely truncated 
apices bearing a short lateral tooth and a long, curved, distal tooth. 
4, stonei, new species 


Parameres broadest at bases or without truncated apices____________ 6 
Stems of parameres very slender and rodlike___-_________________-- a 
Stems of parameres bladelike! 2)... 22 L ee eee 9 


Apices of parameres slightly expanded, palplike; aedeagal sclerites 
sinuate, stouter toward apices; ninth sternite with deep, quadrate, 
mesal notch; dististyles very broad, with truncate apices. 

1. elegantula (Johannsen) 

Apices of parameres sharp-pointed and bent ventrad; aedeagal sclerites 
bént more than twicess 252 22 le ee eee 8 

Membrane of ninth sternite bare; tips of aedeagal sclerites recurved 
hooklike; dististyles very slender__.__._-__- 6. beckae new species 

Membrane spiculate; tips of aedeagel sclerites snoutlike and crossing 
each other; dististyles stout_....__----- 14, punctipes, new species 

Parameres with an extra lateral pair of straight, bladelike arms half 
as long as inner arms; membrane of ninth sternite bare. 

15. rabelloi Lane 

Parameres simple, without lateral arms, expanded subapically, with tips 
abruptly narrowed; membrane spiculate________ 3. lutea (Malloch) 

Parameres with knobbed apices over twice as broad as the very slender, 
stalklike stems; basistyles very large and globular; dististyle very 


slender “to bases 42 PUN eu: See eee 18. viridis (Coquillett) 
Parameres otherwise, without knobbed apices____._..._.._------- 11 
Parameres with greatly swollen, conical bases and very slender, tapered 
HpPICes.. wei ete chee estos tt NS OS eee 12 
Parameres with stems of subequal diameter or swollen towards 
Apiceseick 222 St sees bela eke elec eee 13 
Parameres with basal 0.6 swollen, the pointed apices abruptly bent 
towards base on outer side______________- 10. diversa (Coquillett) 


Parameres with only basal 0.2 swollen, distal portions consisting of very 
slender, curved, saber-shaped blades crossing each other in middle. 
16. sybleae, new species 

Parameres nearly straight, with apices abruptly narrowed and bent 


OGD... 32Sus: | OSE 2s oe ee eee ee 14 
Parameres sinuate or with broad, toothlike, lateral, subapical expan- 
SIONS: 22 Ub 522 Jo 5 ee De een ee ee 15 
Parameres with apices bent outward__----_--------- 7. bicolor Lane 
Parameres with apices bent inward___--------_--- 11. glauca Macfie 


Lobe of basistyle quadrate, as long as broad; parameres slender and 
markedly sinuate; dististyles greatly curved. 

5. antennalis (Coquillett) 

Lobe of basistyle consisting of only a hump on inner margin; parameres 

very broad and stout; dististyle nearly straight_____._._________- 16 

Each paramere with broad, toothlike, lateral, subapical expansion about 

half as broad as length of dististyle, with a slender, ventrally curved 
spine from apex on inner margin; dististyle tapered. 

9. coquilletti Kieffer 

Each paramere swollen slightly midway to apex without subapical lateral 

expansion; dististyle with broad, blunt apex... 13. pruinosa Wirth 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 61 


TaBLeE 1.—Proportions of segments of hind legs of female Stilobezzia species. 


Species Cxo Vir F tr 4 Ts T3 Ty T; 

ee ee —_|————_ 

eelerantulans <4: 2eose in dela 20 | 10 | 60 |} 60 | 30 | 10 5 4 12 
a a3 Ait 25 | 10 | 60 | 65 | 32 | 15 6 4 8 
ee 40 | 15 | 90 | 90 | 40} 20] 9 6 15 
DOMME ie eo AO VOL ao | BO LS fa SD 10 
Pmatuonmeise so fo TE SEE ZO 12 rod fod |" 28° | 12 3 3 8 
BamperAel tl: OF9)b eo LLCS. L380) Pb |) 75) 7S) fe 40 | 18 6 5 8 
TSS es a peel lb 7 | 40{ 35 | 17) 10 + 3 10 
Pee so eH opel es 15 hole Bone eee kb 8; 3 2 5 
Peomuilertt._.--_.._.-.......| 80 | 15 | 80 | 85 | 40 | 18 5 5 13 
LOM cth( | ——— ee 8 6 = 5, 13 6] 35 | 82] 16 6 3 3 5 
eeaetenea | 20 10 | 65-| 70 | 33 15 4| 4 10 
ESSELTE SRS Sa SS es eee See eee (ee ee ee ee 
2 20 | 10; 50} 50} 28 | 10| 4] 4 8 
MEIPPICUINOS= = = 2 _.| 30 | 15 | 80 | 80 | 40 | 20 6 4 10 
A Shih 5 nie 40 |, 15)| 85.) 95 | 45.| 25.) 10 5 15 
0 SE re 15 @ 45.) 50.) 25 | 12 3 2 5 
OL 20 | 10} 55 | 55 | 28 | 12 3 3 8 


Subgenus Eukraiohelea Ingram and Macfie, 1921 


Eukraiohelea Ingram and Macfie, Ann. Trop. Med. Parasit., vol. 15, p. 347, 1921.— 
Johannsen, Arch. Hydrobiol., Suppl., vol. 9, p. 430, 1931; Journ. New York 
Ent. Soc., vol. 42, p. 344, 1934.—Macfie, Ann. Trop. Med. Parasit., vol. 34, 
p. 22, 1940.—Tokunaga, Tenthredo, vol. 3, p. 344, 1940.—Johannsen, Ann. 
Ent. Soc. Amer., vol. 36, p. 781, 1943; Bull. Bishop Mus. No. 189, p. 190, 1946. 
(Genotype, Hukraiohelea africana Ingram and Macfie, 1921; designated by 
Macfie, 1940.) 


Diagnosis.—First radial cell absent, the r-m crossvein nearly form- 
ing a straight line with Ri, the latter arising just a little more toward 
base of wing; fore femora with 2 or 3 ventral spines but not swollen; 
male aedeagus with hyaline posterior membrane, parameres long and 
very slender. 

Discussion.—Johannsen (1934) placed Eukraiohelea as a subgenus 
of Parabezzia Malloch, 1915. However, as shown by a comparison 
of the male genitalia of P. petiolata Malloch, 1915, with those of the 
four species of Hukraiohelea whose males have been described, the 
latter are much more closely related to Stilobezzia. The following 
species have been correctly placed in Eukraiohelea: E. africana Ingram 
and Macfie, 1921, #. versicolor Ingram and Macfie, 1921, and EL. foyz 
Ingram and Macfie, 1922, all from West Africa; Stilobezzia (Hukraio- 
helea) aberrans Johannsen, 1931, from Java; S. (E.) esakiana Tokunaga, 
1940, from the Caroline Islands; and Hukraiohelea amnigena Macfie, 
1935, from Brazil. Hukraiohelea inusitata Johannsen, 1946, from 


62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


Guam is apparently a synonym of esakiana Tokunaga. Palpomyia 
dorsofasciata Lutz, 1914, from Brazil was correctly referred to Hukraio- 
helea by Macfie (1935). Parabezzia poikiloptera Ingram and Macfie, 
1922, from West Africa should be referred to Hukraiohelea as tenta- 
tively suggested by Johannsen (1946), but the position of S. ugandae 
Ingram and Macfie, 1923, from East Africa, which Johannsen also 
believed to belong here, is more doubtful. De Meillon (1938), more- 
over, states that versicolor and poikiloptera should be placed in the 
subgenus Stilobezzia, since material he determined as these two species 
from Lourengo Marques, Mozambique, possessed a small but distinct 
first radial cell. 


1. Stilobezzia (Eukraiohelea) elegantula (Johannsen), 1908, new combination 
Fieure 11, g 


Bezzia elegantulus Johannsen, Kansas Univ. Sci. Bull., vol. 4, p. 109, 1907 (9, 
Kansas). 

Probezzia elegantula, Malloch, Proc. Biol. Soc. Washington, vol. 27, p. 137, 1914. 

Parabezzia elegantula, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 359, 
1915.—Johannsen, Journ. New York Ent. Soc., vol. 42, p. 345, 1934. 

Eukraiohelea elegantula, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. 


Description. —FEMALE: Length 2.0 mm., wing 1.5 mm. by 0.7 mm. 
A yellowish species, some specimens with a greenish tinge; antennae 
brown, pedicel and narrow bases of flagellar segments yellow; palpi 
dark brown. Mesonotum deep yellow, with broad sublateral brown 
bands; scutellum yellow in center, ends brown; postscutellum brown; 
pleura grayish, a large black spot on mesopleuron. Legs yellow; coxae 
dark below; trochanters brown; apex of hind femur, base of hind tibia, 
apices of all tibiae and narrow apices of all tarsal segments brown. 
Wings, including veins, grayish hyaline; halteres yellow, sides of knobs 
black, flat end of knobs pale green. Abdomen dark grayish brown, 
apices of tergites with narrow white bands. 

Antennae very long and slender. Mesonotum with long black 
hairs in rows; scutellum with four long black marginal bristles and a 
few short hairs. Legs with hairs fine except on hind tibia where they 
are long and black; proportions of segments of hind leg as in table 1; 
fore femur with two or three small black spines midway on inner 
margin; basitarsus with a long black basal and a smaller distal spine 
on midlegs and hind legs; fifth tarsal segment with two pairs of long 
black ventral batonnets; claws slender and unequal, the outer claw 
as long as fifth segment, the other about a third as long. Wing with- 
out macrotrichiae; first radial cell absent, r-m crossvein and R, nearly 
forming an oblique line, lengths of the two sections of costa in ratio 
of 0:6; petiole of media as long as r-m crossvein. 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 63 


Maus: Ninth sternite nearly four times as broad as long, with deep 
quadrate emargination three-fourths way to base on mesal third, with 
spiculate margins, the posterior membrane bare; ninth tergite rounded, 
with a pair of very small fingerlike setigerous apicolateral lobes. 
Basistyles simple, about twice as long as broad; dististyles about two- 
thirds as long as basistyles, very broad with truncate tips. Aedeagus 
with an oblique pair of rather stout, slightly sinuate, sclerotized bars 
with pointed apices; an irregular hyaline membranous lobe from pos- 
terior margins. Parameres very long, slender and rodlike, with slender 
lateral apodemes; stems about a fourth again as long as basistyles, 
subparallel, the apices slightly swollen, palplike, and bent ventro- 
laterad. 

Type.—In Snow collection, University of Kansas, 9, Kansas, 
Douglas County, Lawrence. 

Material examined.—Florida: Crystal River, Citrus County, July 
7, 1949, Sept. 18, 1950, Hudson, 2 99, Miami, Dade County, Oct. 15, 
1947, Buren, 1 9; Panama City Beach, Bay County, July 2, 1950, 
McElvey, 1 0 

Louisiana: Baton Rouge, Apr. 30, 1947, Wirth, 1 9. 

Jamaica: Newport, Feb. 22, 1937, Chapin and Blackwelder, 1 9. 

Remarks.—S. (E.) africana Ingram and Macfie, from West Africa, 
the genotype of Eukraiohelea, is almost identical with elegantula, but 
is smaller and the male genitalia differ markedly; the ninth sternite 
does not have the mesal excavation, the dististyles have the apices 
slenderer and rounded, the apices of the parameres are straight and 
pointed, and the membrane of the aedeagus is prolonged mesad in a 
sharp cone. 


Subgenus Neostilobezaia Goetghebuer 


Neostilobezzia Goetghebuer, in Lindner, Die Fliegen der Palaearktischen Region, 
Lief. 78, p. 53, 1934. (Genotype, Ceratopogon ochraceus Winnertz, 1852, 
by present designation.) 


Discussion.—Goetghebuer (1934) distinguished this subgenus from 
Stilobezzia by the presence of macrotrichiae at the end of the wing, 
the wing of Stilobezzia being entirely bare. In addition, all the species 
known to me are yellowish or brown in ground color, varying from 
almost whitish to almost black, and the male genitalia have the 
aedeagal sclerites decidedly bent or humped in the middle and the 
basistyle lacks the inner Jobe. Species with these characters are 
widely distributed around the world. It is especially significant that 
most of the species described from the temperate regions belong to 
this subgenus, while the tropical species are predominantly the sub- 
genus Stilobezzia. 


64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


2. Stilobezzia (Neostilobezzia) fuscula Wirth, 1952 
Ficure 11, e 


Stilobezzia fuscula Wirth, Univ. California Publ. Ent., vol. 9, p. 204, 1952 (9, 
California). 


Diagnostic characters —This species resembles S. (NV). lutea (Mal- 
loch), 1918, in its uniformly yellowish color, unmarked wings with 
macrotrichiae, and mesonotum with indistinctly darker vittae; but 
differs in being somewhat duller and darker, with dark antennal 
pedicel and halteres, and in having the body hairs much softer. 

Description —F rmaE: Length 1.4 mm., wing 1.5 mm. by 0.5 mm. 
Head grayish brown pruinose, pedicel of antennae and palpi brown, 
antennal flagellum yellowish brown; segments in proportion of 
20:12:12:12:12:12:12:12:20:20:20:20:30. Palpal segments in pro- 
portion of 4:8:15:10:10, third segment with three long, stalked sen- 
sillae. 

Mesonotum dull yellowish brown, with coarse pruinosity, a narrow 
median longitudinal vitta and a broad patch on each side from humeral 
pits to wing base darker brown; most of bristles removed, but those 
few remaining long and dark brown. Scutellum bright yellow in 
middle, brownish on sides, apparently with four marginal bristles. 
Postscutellum and pleura yellowish brown pruinose. Legs dull dark 
yellowish; hairs short, except row of long bristles on outer edge of hind 
tibia; inner claw as long as fifth tarsal segment. 

Wings grayish hyaline, anterior veins brownish; second anterior 
radial cell about five times as long as first; macrotrichiae thick at apex 
of cell R; and along and beyond intercalary fork, a few also along tip 
of veins M, and M, and at apex of cell M;. Halteres brown, flat end of 
knob dull yellowish. 

Abdomen dull dark brown, a patch of long brown hairs on sides 
of first segment. Spermathecae two, oval, with ducts sclerotized 
a short distance; a rudimentary third spermatheca present. 

Mate: Similar to the female but with the usual sexual differences; 
antennal plumes light brown; thorax and abdomen infuscated light 
brown; scutellum yellow. 

Ninth sternite about three times as broad as long, with mesal 
posterior excavation halfway to base, posterior membrane bare; ninth 
tergite rather long and rounded, with a pair of long, rounded, seti- 
gerous, apicolateral lobes. Basistyle nearly three times as long as 
broad, with a black spine borne on a small lobe with a long fine hair 
about midway of mesal margin; dististyle as long as basistyle, slender 
and slightly curved to apex. Aedeagus with a pair of oblique, lateral, 
sclerotized bars with median apices forked, the inner arm in line 
with base and longer than the other arm. Parameres with slender 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 65 


basal apodemes, stems long and slender, with bases slightly swollen, 
each with apex gradually narrowed to a fine apical filament abruptly 
bent and recurved on distal fourth. 

Type-—USNM 59946, 9, California, Tulare County, 6 miles east of 
Orosi. (In the original publication the sex was erroneously given 
as o’.) 

Material examined.—California: 6 miles east of Orosi, Tulare 
County, July 8, 1947, Wirth, 2 99 (type and paratype). 

Utah: Moab, June 8, 1948, Knowlton, Harmston, and Wood, at 
light, 1 o@. 

Remarks.—The male genitalia of S. fuscula are practically identical 
with those of S. macfiei Lane, 1947, from Brazil. However, the latter 
species has the mesonotum polished black, with lighter areas, and 
under certain light a spot on each side before wing base has purplish 
reflections. Moreover, the absence of macrotrichiae on the wings 
would place S. macfiei in the subgenus Stilobezzia. 


3. Stilobezzia (Neostilobezzia) lutea (Malloch), 1918 
FicureE 11, 6 


Hartomyia gilua Malloch, not Coquillett (misidentification), Bull. Illinois State 
Lab. Nat. Hist., vol. 10, p. 343, 1915. 

Hartomyia lutea Malloch, Bull. Brooklyn Ent. Soe., vol. 13, p. 18, 1918, (¢ , Illinois). 

Stilobezzia lutea, Johannsen, Ann. Ent. Soc. Amer. vol. 36, p. 781, 1943. 

Stilobezzia mallochi Hoffman, Ent. News, vol. 35, p. 283, 1924, (o", 9; New York, 
Pennsylvania).—Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943 
(eastern United States). (New synonymy.) 

Description —FrEMA.LE: Length 2.5 mm., wing 1.9 mm. by 0.7 mm. 
Yellow, including anterior wing veins, antennae, and legs; mesonotum, 
except humeri, and apex of abdomen rufous brown. Vestiture of 
very long coarse brown bristles, those on scutellum as long as breadth 
of scutellum. 

Mats: Ninth sternite nearly three times as broad as long, with a 
very broad, shallow, posterior emargination, the membrane spiculate; 
ninth tergite conical, with a prominent, submedian pair of long, fleshy, 
setose lobes. Basistyles simple, twice as long as broad; dististyles 
slightly longer than basistyles, curved, and gradually attenuated, 
with pointed apices. Aedeagus with a pair of oblique sinuate 
sclerites, bases of which are articulated with ventral root of basistyle, 
apices close together, pointed, and bent ventrad. Parameres with 
stout basal apodemes, the stems close together and slightly sinuate, 
flattened and slightly expanded past middle, with apices pointed and 
somewhat curved ventrad. 

Type.—tin collection Illinois Natural History Survey, 2, Illinois, 
Elizabeth. 

Material examined—New Hampshire: Center Harbor, July 9, 
Dyar, 1 9; Franconia, Slosson, 1 o. 

232991532 


66 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 103 


Connecticut: Green Falls, June 31, 1935, Chapman, 1 &, 1 9. 

Rhode Island: Westerly, June 10, 1936, Chapman, 1 <. 

New York: Armonk, Bemus Point, Canadarago Lake, Kast 
Bridge, Millwood, Poughkeepsie, Rome, June, July 1934-36, H. K. 
Townes, 12 oo’, 7 99. 

New Jersey: Moorestown, June 6, 1936, H. and M. Townes, 1 9; 
Riverside, June 18, 1939, H. K. Townes, 1 &. 

Delaware: Smyrna, June 8, 1937, Bradley, 1 &. 

Maryland: Cabin John, June 18, 1920, Aldrich, 1 2; Glen Echo, 
July 1, 1923, Malloch, 19; Plummer’s Island, June 18, 1914, Shannon, 
i. 9. 

District of Columbia: Rock Creek Park, June 17, 1920, Aldrich, 10. 

Virginia: Dead Run, June 6, 1914, Shannon, 1 9; Falls Church, 
June 7, 1914, Shannon, 2 99, and July 4, 8, 1950, Wirth, 4 oo, 7 99; 
Glencarlyn, June 7, 1935, Malloch, 1 @, 1 9. 

South Carolina: Greenville, June 4, 1933, H. K. Townes, 1 &. 

Indiana: Lafayette, June 26, 1916, Aldrich, 1 o, 4 99. 

Illinois: Elizabeth, July 7, 1917, Malloch, 1 9 (type of Hartomyia 
lutea Malloch). 

Michigan: Cheboygan County, July 17, 1942, Sabrosky, 1 9. 

Iowa: Maquoka Caves State Park, Jackson County, July 3, 1949, 
Laffoon, 1 9; Pikes Peak State Park, Clayton County, July 4, 1949, 
Laffoon, 1 9; Sioux City, June, July 1949-50, Slater and Laffoon, 
6 uct 199: 

Remarks.—Malloch had misidentified the male of this species as 
Hartomyia gilva (Coquillett), which is an Atrichopogon, in 1915, and 
in 1918 described the female of the same species as Hartomyia lutea, 
new species. Hoffman discovered that Malloch’s determination of 
gulva was incorrect and described the species as Siilohezzia mallochi, 
1924, in part from Malloch’s material. Since Hoffman made no 
mention of Malloch’s description of /utea, he may have been unaware 
of it, or he may not have been able to recognize Malloch’s species. 
The present synonymy has been made after an examination of the 
type of lutea and specimens determined as gilva by Malloch in the 
Ulinois Natural History Survey through the kindness of Dr. H. H. 
Ross, and of specimens in the U. 8. National Museum determined by 
Hoffman as mallochi. Hoffman’s detailed description of this species 
needs no elaboration except for the characters of the male genitalia. 


4. Stilobezzia (Neostilobezzia) stonei, new species 
Figure 11, c 


Diagnostic characters.—A large brownish species; mesonotum and 
scutellum dull yellowish; legs yellowish; midfemora, hind femora, and 
tibiae dark brown; wings grayish brown; halteres dull yellow. 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 67 


Description.—Frma.e: Length 1.8 mm., wing 1.8 mm. by 0.7 mm. 
Head dull brownish, including antennae and palpi, with a few long 
black hairs. Antenna as long as head and thorax combined; flagellar 
segments in proportion of 20:15:15:15:15:16:17:18:25:25:25:25:30. 
Palpal segments slender, in proportion of 5:10:12:6:10, third segment 
slightly swollen on basal two-thirds, with a small pit at distal fourth. 

Thorax dull brown, mesonotum and scutellum dull yellowish, 
postscutellum and pleura dark brown. Mesonotum with moderate 
to long brown hairs in rows; scutellum with about ten long brown 
marginal hairs. Legs dull brown, usually all of forelegs, midtarsus 
and hind tarsus yellowish. Proportions of segments of hind leg as in 
table 1; legs moderately slender, hairs fine, not very long except 
those on hind tibia about twice the diameter of segment; basitarsi 
with ventral spines, three or four on basal half on forelegs and midlegs, 
one at base on hind leg; claws unequal, rather strong and curved, the 
longer nearly as long as fifth segment, the other about half as long. 

Wings evenly infuscated, the veins slightly darker, a few macro- 
trichiae at apices of cells R; and M,. First radial cell large, about a 
third the length of second; costa extending to 0.8 wing length; cross- 
vein r-m faint in middle, oblique; petiole of media as long as crossvein. 
Halteres yellow. Abdomen dark brown with short fine hairs; two 
large, slightly unequal ovoid spermathecae and a vestigial third 
present, ducts not sclerotized. 

Mats: Similar to the female but with the usual sexual differences; 
antennal plumes brownish, knees quite dark. Ninth sternite about 
three times as broad as long, with broad posterior excavation more 
than halfway to base, the membrane bare; ninth tergite rounded, with 
small, submedian, setose, apicolateral lobes. Basistyles simple, twice 
as long as broad; dististyles 0.9 times as long as basistyles, slender and 
nearly straight, their apices slightly knobbed. Aedeagus with an 
oblique pair of slightly curved sclerites with distal ends close together, 
sharp-pointed, and bent ventrad; apparently bearing a pair of hyaline 
membranous lobes on posterior margins and attached to membrane 
connecting dorsal roots of basistyles on ventral side of parameres. 
Parameres with broad, platelike, basal apodemes; stems very broad 
and flattened, gradually expanded distad with a short lateral point and 
a longer, slenderer, ventrally bent, distal point, their distal margins 
obliquely truncated. 

Types —USNM 60964, holotype, &, and allotype, Falls Church, 
Va., July 8, 1950, Wirth. Paratypes: Virginia: 16 io, 24 99, same 
data as type. Georgia:2 oo, 17 99, Thomasville, May 1949, Palmer. 
Florida: 2 99, Innerarity Point, May 4, 1950, Rathert; 1 9, Panama 
City Beach, May 6, 1949, McElvey. 


68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Remarks.—Related to S. lutea (Malloch) in general features, but 
readily distinguished by the more extensive brown coloration, 
finer vestiture and the characteristic broad, obliquely truncated, two- 
pointed, platelike, male parameres. This species is named in honor of 
Dr. Alan Stone. 


Subgenus Stilob ezzia Kieffer, 1911 


Stilobezzia Kieffer, Rec. Indian Mus., vol. 6, p. 118, 1911, and vol. 9, p. 184, 1913; 
Bull. Ent. Soc. France, p. 192, 1919.—Goetghebuer, Mem. Mus. Hist. Nat. 
Belgique, vol. 8, p. 59, 1920.—Carter, Ingram, and Macfie, Ann. Trop. Med. 
Parasit., vol. 15, p. 324, 1921.—Edwards, Trans. Ent. Soc. London, vol. 74, 
p. 411, 1926; Notulae Ent., vol. 9, p. 9, 1929.—Johannsen, Arch. Hydrobiol., 
Suppl., vol. 9, p. 430, 1931; Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943.— 
Ingram and Macfie, The Diptera of Patagonia and southern Chile . . ., 
pt. 2, fase. 4, p. 191, 1931.—de Meillon, Proc. Ent. Soc. London, ser. B, vol. 
7, p. 266.—Tokunaga, Philippine Journ. Sci., vol. 72, p. 155, 1940.—Lane, 
Rev. Ent., vol. 18, p. 197, 1947.—Lee, Proc. Linn. Soc. New South Wales, 
vol. 72, p. 345, 1948.— Wirth, Univ. California Publ. Ent., vol. 9, p. 202, 1952. 
(Genotype, Ceratopogon notatus de Meijere, as Stilobezzia festiva Kieffer; 
original designation.) 

Hartomyia Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 339, 1915. 
(Genotype, Ceratopogon pictus Coquillett; original designation.) 


Diagnosis.—Two radial cells present; wing without macrotrichiae 
at apex. 


5. Stilobezzia (Stilobezzia) antennalis (Coquillett), 1901 
Ficure 12, e 


Ceratopogon antennalis Coquillett, Proc. U. 8S. Nat. Mus., vol. 23, p. 606, 1901 
(2, District of Columbia). 
Ceratolophus antennalis, Kieffer, Genera insectorum, fasc. 42, p. 60, 1906. 
Johannseniella antennalis, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, 
p. 227, 1914. 
Hartomyia antennalis, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 348, 
1915 (o; Illinois, Indiana). 
Stilobezzia antennalis, Johannsen, Ann. Ent. Soc. Amer., 36: 781, 1948. 
Description. FEMALE: Length 1.5 mm., wing 1.6 mm. by 0.6 mm. 
Head, thorax, and abdomen polished black; antennae and legs yellow, 
midfemur and hind femur black except at extreme bases. Wings bare 
and hyaline, anterior veins yellowish, knob of halteres black. Palpi 
and last five flagellar segments slightly infuscated, flagellar segments 
in proportion of 18:11:211:11:11:11:11:15:25:25:25:25.20)" Pa 
segments in proportion of 4:8:14:8:15, third segment not swollen, 
pit absent. Proportions of segments of hind leg as in table 1; basitarsi 
without spines, fifth segment with a pair of long ventral batonnets; 
claws long, simple, and very unequal, the short claw fused with base 
of and about a third as long as the other. First radial cell about a 
fourth as Jong as second; r-m crossvein vertical, petiole of media to a 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 69 


third the length of M;. Spermathecae two, the larger oval, the 
smaller subspherical and a third the diameter of other, a minute 
vestigial third present. 

Mate: Similar to the female but with the usual sexual differences; 
plumes of antennae brown. Ninth sternite a narrow transverse ante- 
rior band, the membrane spiculate; ninth tergite evenly rounded 
caudad, the apicolateral lobes not prominent. Basistyle with large 
irregular lobe on inner side to distal third; dististyle rather strongly 
bent past middle and tapered to pointed tip. Aedeagus with a pair 
of oblique, straight, sclerotized bars with distal apices scarcely meet- 
ing. Parameres a pair of strongly sclerotized, large, strongly sinuate 
rods with pointed apices and large basal knobs. 

Type.—USNM. 5481, 9, District of Columbia. 

Material examined—New York: Ithaca, June 11, 1935, H. K. 
Townes, 1 9; Rome, June 24, 1934, H. K. Townes, 1 &@. 

New Jersey: Moorestown, June 6, 1939, H. and M. Townes, 1 o. 

Maryland: Dorchester, July 10, 1907, Barber, 1 9; Mayo, July 22, 
1950, Wirth, 2 <7, 1 9; Plummer’s Island, June 30, 1914, Shannon, 
1 9, and Aug. 18, 1913, Viereck, 1 @. 

District of Columbia: June, 1 9 (type); Benning’s, July 21, Aldrich, 
2 99. 

Virginia: June 16, 1903, Pergande, 2 7, 3 29; Dead Run, June 
22, 1915, Shannon, 1 9, Falls Church, July 4, 8, 1950, Wirth, 3 #7, 
4 99. 

Michigan: Midland County, July 17, 1944, Dreisbach, 1 2; Traverse 
City, June 17, 1943, Sabrosky, 1 @. 

Illinois: Freeport, July 4, 1917, Malloch, 3 oo; Galena Junction, 
July 8, 1917, Malloch, 1 9; White Heath, June 4, 1939, Dirks, 1 9. 

Tennessee: Knoxville, May 25, Aldrich, 1 9. 

Mississippi: West Point, Aug. 11, 1904, Barber, 1 @. 

Louisiana: Baton Rouge, May 4, 16, 1947, Wirth, 4 #0, 8 99. 


6. Stilobezzia (Stilobezzia) beckae, new species 
Fiaure 11, f 


Diagnostic characters.—A large, dark brown species with silvery 
pruinose pattern on mesonotum and abdomen; wing with two darker 
and one lighter spots on anterior margin; knobs of halteres black; 
mesonotum with strong anterior tubercle; fifth tarsal segment 
unspined. 

Description.—FrEmMaueE: Length 2.5 mm., wing 1.8 mm. by 0.7 mm. 
Head black with dense whitish pruinosity, antennae brown, bases of 
flagellar segments 3 to 10 yellowish; palpi dark brown. Flagellar 
segments of antenna in proportion of 16:12:15:20:20:20:20:22:28: 


70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


30:32:32:42. Palpal segments in proportion of 5:12:15:8:12, 
third segment slightly swollen, with large, shallow sensory pit near 
apex. 

Thorax dark reddish brown, with dense silvery pruinescence 1 
irregular patches; mesonotum long, with prominent, conical, anterior 
spine, the anterior portion markedly sloping, posterior portion flat- 
tened between wing bases; covered with sparse, long, brown hairs; 
scutellum with four long, fine, brown hairs. Legs dark brown, all 
femora and fore tibiae with obscure broad median light bands, mid- 
tibia and hind tibia with narrow sub-basal light rings, tarsi yellow 
except fifth segment of each brown. Proportions of segments of 
hind leg as in table 1; legs with moderately long, black hairs, which are 
stronger on hind tibia; basitarsi with three ventral spines on foreleg, 
four on midleg, and one at base on hind leg; fifth segment without 
ventral batonnets; claws very slender and slightly unequal, the longer 
as long as fifth segment, the other about 0.6 to 0.8 as long. 

Wing hyaline, without macrotrichiae, veins infuscated, a large 
brownish spot over first radial cell from vein R, to crossvein r-m; 
apex of second radial cell with brown spot, then a short white area 
with apex of wing weakly infuscated. First radial cell well developed, 
one-fourth as long as second; r-m crossvein faint in middle, forming an 
oblique line with R,, petiole of media as long as crossvein. Knobs of 
halteres black. 

Abdomen black, distal half of tergites 1 to 3 and narrow distal 
margins of remaining tergites with silvery, pruimose bands. Sper- 
mathecae two, subequal, pyriform. 

Mats: Similar to the female but with the usual sexual differences; 
antennal plumes golden. Ninth sternite over twice as broad as long, 
with caudal excavation to half its length, the membrane bare; ninth 
tergite rounded with a pair of membranous, setigerous, apicolateral 
lobes. Basistyles simple, about 2.5 times as long as broad; dististyles 
slender, nearly as long as basistyles, gently curved, with pointed 
incurved apices. Aedeagus with a pair of heavy, oblique, crooked, 
lateral sclerites meeting mesad subapically, with apices pointed and 
abruptly bent ventrolaterad. Parameres with heavy, platelike, 
V-shaped, basal apodemes; stems straight and contiguous mesad, 
slightly expanded midway, with lateral wall folded ventrad and mesad 
and apparently fusing on midline on distal third; extreme apices 
slender, flattened dorsoventrally and abruptly bent ventrad. 

Types —USNM 60695, holotype, , and allotype, Crystal River, 
Citrus County, Fla., June 7, 1950, Hudson, light trap. Paratypes: 
Florida: 10 o'o7,17 99, same data as type except dates, June 5—July 17, 
1949-50; 6 99, Grayton Beach, Walton County, May 10, 1949, Butler; 
15 99, Santa Rosa, Walton County, June 7, 1949, Peterson. Missis- 
sippi: 4 99, Horn Island, June, July, 1944, Richmond. 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH va 


Additional material examined.—Pert: Iquitos, March, April 1931, 
Shannon, 1 9. 

Remarks.—Closely allied to S. paulistensis Lane, 1947, from Brazil 
which differs, however, in having the legs yellowish rather than black- 
ened, in lacking the dark spot at apex of second radial cell, and in 
having the lateral sclerites of the male aedeagus shorter, with apices 
capitate and bent mesad rather than slender and bent laterad. S. 
rabellot Lane, 1947, is also closely related, but is a larger, hairier 
species with three dark wing spots, including one at apex of first 
radial cell, and has several pairs of batonnets on fifth tarsal segment, 
IT am happy to name this species for Mrs. Elisabeth C. Beck of Arling- 
ton, Fla., who has kindly sent me so many interesting Florida heleids, 


7. Stilobezzia (Stilobezzia) bicolor Lane, 1947 


Ficure 11, 7 


Stilobezzia bicolor Lane, Rev. Ent., vol. 18, p. 208, 1947 (o, Brazil). 


Description.—F mMALE: (Here described for the first time.) Length 
1.5 mm., wing 1.5 mm. by 0.6 mm. Head dark brown, antennal 
pedicel, clypeus, and palpi yellowish. Thorax and abdomen polished 
black, with metallic bluish violet reflections. Mesonotum rather 
broad and flat, with sparse, long, brown hairs, scutellum with four 
marginal bristles. Midcoxa and hind coxa black, rest of legs yellow, 
except knees and fifth tarsal segments dark. Legs with fine hairs; 
proportions of segments as in table 1; basitarsi unspined; fifth segment 
with a pair of long black batonnets at base: claws slender and unequal, 
the outer claw as long as fifth segment, the inner about a third as long. 
Wing hyaline, with violet reflections, anterior veins yellow; macrotri- 
chiae absent. First radial cell distinct, a sixth as long as second; 
petiole of media about twice as long as crossveinr-m. Halteres black. 
Abdomen convex above, somewhat petiolate, with a row of long, brown 
hairs across each tergite. 

Mate: As in the female but with the usual sexual differences; 
plumes of antennae brown. Ninth sternite a narrow anterior band, 
the posterior membrane spiculate; ninth tergite rounded, with incon- 
spicuous, rounded, setigerous, apicolateral lobes. Basistyle with a 
small lobe at half the length of inner margin; dististyle about as long 
as basistyle, tapered to slender tip. Aedeagus with an only slightly 
oblique pair of sclerotized bars, stout in the Texas specimen figured, 
but slenderer in the tropical specimens. Parameres with lateral 
apodemes bent caudad; stems straight, stout and rodlike, with beak- 
like apices abruptly bent laterad. 

Type.—In collection University of Sao Paulo, Brazil, @, Brazil, 
Sao Paulo, Osasco. 


72 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


See 


c. STONE! 


. FUSCULA ai 
: f- BECKAE 


d. PUNCTIPES 


g- ELEGANTULA h. COQUILLETTI i. BICOLOR 


Ficure 11.—Male genitalia of Stilobezzia species: a, S. (S.) rabelloi Lane; b, S. (N.) lutea 
(Malloch); c. S. (N.) stonei, new species; d, S. (S.) punctipes, new species; ¢, S. (N.) 
fuscula Wirth; f, S. (S.) beckae, new species; g, S. (£.) elegantula (Johannsen); h, S. (S.) 
coquilletti Kieffer; 1, S. (S.) bicolor Lane. 


HELEID MIDGES, GENUS STILOBEZZIAWIRTH 73 


d, VIRIDIS e. ANTENNALIS 


f. DIVERSA g. PRUINOSA h. GLAUCA 


Ficure 12.—Male genitalia of Stilobezzia species: a, S. (S.) thomsenae, new species; b, S. 
(S.) bulla Thomsen; c, S. (S.) sybleae, new species; d, S. (S.) viridis (Coquillett); ¢, S. (S.) 
antennalis (Coquillett); f, S. (S.) diversa (Coquillett); g, S. (S.) pruinosa Wirth; h, S. 
(S.) glauca Macfie. 


74 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Material ecamined.—Texas: San Antonio, Apr. 8, 1907, Pratt, 1 @. 

Costa Rica: Higuito, Schild, 2 7, 10 99. 

Puerto Rico: Anasco, Apr. 22, 1933, Faxon, Mills, and Anderson, 
1 &#, 499. Bayamon, Jan. 28, 1934, Lesesner and Anderson, 1 9. 

Panama: Summit, Canal Zone, November 1946, Krauss, 2 o'o7,19. 


8. Stilobezzia (Stilobezzia) bulla Thomsen, 1935 


Figure 12, 6 


Stilobezzia bulla Thomsen, Journ. New York Ent. Soc., vol. 438, p. 289, 1935 
(o, 2; New York); Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. 


Description FEMALE. Length 1.0 mm., wing 0.8 mm. by 0.4 mm. 
Pruinose grayish brown; scutellum and legs yellowish; halteres white; 
wings clear, veins stramineous. Antennal segments in proportion of 
12:8:9:10:10:10:10:10:13:15:15:15:20. Palpi short and slender, 
segments in proportion of 5:7:7:5:7, third segment not swollen, pit 
absent. Basitarsi and fifth segment without spines; claws simple, 
slender and unequal, the longer as long as fifth segment, the other 
a third as long. Wings clear, macrotrichiae absent. Spermathecae 
two, oval, slightly unequal, with ducts sclerotized a very short 
distance. 

Maus: Ninth sternite a narrow anterior ribbon, the membrane 
spiculate; ninth tergite tapered to a pair of prominent, slender, setose, 
apicolateral lobes, truncate between their bases. Basistyle slender, 
with a prominent, heavily sclerotized, platelike lobe articulating with 
the median sclerite (? of aedeagus); dististyle about half as long as 
basistyle, rather stout and nearly straight, tip bluntly pointed, with 
a distinct subapical tooth on ventral face of outer edge. Aedeagus 
with basal sclerites reduced to very small, narrow bars, connected at 
right angles by a pair of short, barlike sclerites to a more dorsal, large, 
heavily sclerotized, transverse sclerite shaped like a pair of buffalo 
horns, with points articulating with inner lobes of basistyles. Para- 
meres with slender, hook-shaped, basal apodemes, stems rodlike, 
slender and nearly straight, gradually tapered to long, simple, fine 
tips reaching nearly to apices of lobes of ninth tergite. 

Types.—In collection Cornell University, holotype, <7, and allo- 
type, New York, Ithaca, McLean Bogs. 

Material examined.—Maryland: Dorchester, July 10, 1907, Barber, 
L ot. 

District of Columbia: Bennings, July 21, Aldrich, 1 9. 

Virginia: Falls Church, July 4, 8, Aug. 6, 1950, Wirth, 18 7d, 
24 99. 

Georgia: Thomasville, May 15-30, 1949, Palmer, 1 9. 

Florida: Crystal River, Sept. 18, 1950, Hudson, 1 9. 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 5 


Louisiana: Baton Rouge, Apr. 19, 1947, Wirth, 1 o&; Kilbourne, 
May 10, 1947, Wirth, 1 9. 


9. Stilobezzia (Stilobezzia) coquilletti Kieffer, 1905 
Ficure 11, h 


Ceratopogon pictus Coquillett, Journ. New York Ent. Soc., vol. 13, p. 60, 1905 
(¢, Virginia). 

Hartomyia picta, Malloch, 1915, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 
341, 1915 (0, Illinois). 

Stilobezzia coquillettt Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 308, 1917 
(new name for C. picta Coquillett not Meigen).—Johannsen, Ann. Ent. Soc. 
Amer., vol. 36, p. 781, 1943. 

Description. —FrmMa.e: Length 2.0 mm., wing 2.0 mm. by 0.8 mm. 
Yellow, in some specimens with greenish tinge; with a pair of widely 
separated, narrow brown lines from anterior median point of meso- 
notum to ends of scutellum; also, narrow center of scutellum, palpi, 
mesopleuron and midcoxa, narrow apices of hind femur and tibia, 
fifth tarsal segment, and fore, lateral, and hind margins of abdominal 
tergites 3 to 7, brown. Wing bare and hyaline, all veins brownish, a 
brown spot over first radial cell including R,; and r-m crossvein; a 
second spot over apex of second radial cell and below halfway across 
cell R;; knobs of halteres brown. 

Flagellar segments of antenna in proportion of 25:15:15:15:15:15: 
17:18:28:28:30:30:60. Palpal segments in proportion of 4:15:20: 
10:22, third segment not swollen, pit absent. Mesonotum and 
abdomen with coarse, dark bristles. Proportions of segments of hind 
leg as in table 1; basitarsi without spines, fifth segment with a pair of 
long, black batonnets at basal third; claws slender and unequal, the 
longer as long as fifth segment, the other about half as long. Sper- 
matheca one, large and ovoid, the duct not sclerotized. Wing with 
macrotrichiae absent; first radial cell very small and narrow, about a 
seventh as long as second, latter to 0.8 wing length; crossvein r-m 
forming an oblique line with vein R,; media petiolate for 0.23 its length. 

Mate: Similar to the female but with the usual sexual differences; 
body more or less greenish with black markings as in female, antennal 
plumes yellow at bases, black on distal halves. Ninth sternite a 
narrow, transverse, anterior band, posterior membrane spiculate; 
ninth tergite full and rounded, with a pair of low, rounded, setigerous, 
apicolateral lobes. Basistyle about twice as long as broad at base, ab- 
ruptly narrowed about midway on inner side, forming alow, mesal lobe 
connected by an irregular membrane to ventrolateral margins of ninth 
tergite; dististyle about 0.7 as long as basistyle, stout, slightly incurved, 
with bluntly pointed tip. Aedeagus with an oblique pair of spatulate 
sclerites with bases abruptly bent laterad around bases of basistyles. 
Parameres a pair of broad, flattened, sclerotized plates, bases connected 
across mid/*ne by a highly sclerotized band from outer anterolateral 


76 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


margins of basistyles; base of each paramere with a prominent lateral 
spur projecting caudad along inner margin of basistyle; distal half 
greatly expanded, with a large, sharp, lateral tooth; inner margins 
contiguous at apices, each with a very slender, pointed spine abruptly 
bent ventrad and cephalad. 

Type—USNM 8356, 9, Virginia. 

Material examined.—Maryland: Shadyside, July 6, 1925, Aldrich, 
19. 

Virginia: June 16, Pergande, 1 2 (type). 

Illinois: Urbana, May 20, 1914, Aldrich, 1 o, 19. 

Louisiana: Baton Rouge, Apr. 13, 25, 1947, Wirth, 3 7d, 3 99. 

México: Ciudad Monte, Tamaulipas, Nov. 23—-Dec. 1, 1943, 
Brookman, 29 oo’, 35 99; Tamazanchale, San Luis Potosi, March 11, 
1944, Brookman, 1 9; Tapachula, Chiapas, Sept. 20, 1944, Brookman, 
ep 

Puerto Rico: Arecibo, Apr. 5, 1932, Faxon and Anderson, 1 o&, 
1 9; San Juan, June 14, 1933, Harlan, 1 9. 

Jamaica: Bath, St. Thomas, Chapin and Blackwelder, 4 992; Spanish 
Town, Feb. 2, 1937, Chapin and Blackwelder, 3 99. 

Panamé: Sabanas, Apr. 20, 1923, Shannon, 1 9. 

Remarks.—S. fiehrigi Kieffer, 1917, from Paraguay and Brazil, 
resembles S. coquilletti very closely except for the abdominal markings, 
with a pair of large lateral brown spots on the first tergite and a large 
median brown spot on the second, and tergites 3 to7 with dark markings 
on the disc rather than on the margins. In the male genitalia of 
fiebrigt the basistyles bear a prominent mesal lobe, the dististyles are 
slenderer at the apices, the lateral bars of the aedeagus are slender 
without capitate tips, and the parameres are not so broad and the 
lateral tooth is not so prominent. 


10. Stilobezzia (Stilobezzia) diversa (Coquillett), 1901 
Figure 12, f 


Ceratopogon diversus Coquillett, Proc. U.S. Nat. Mus., vol. 23, p. 607, 1907 (?, 
New Jersey). 

Ceratolophus diversus, Kieffer, Genera insectorum, fasc. 42, p. 60, 1906. 

Johannseniella diversa, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 


227, 1914. 
Hartomyia diversa, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 344, 


1915. 
Stilobezzia diversa, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. 
Description —Frmaue: Length 1.5 mm., wing 1.4 mm. by 0.6 mm. 
Head brown, antennae and palpi yellow; thorax shining black; abdo- 
men and coxae light green; legs yellow, fifth tarsal segment brownish; 
wings hyaline, anterior veins yellow, halteres yellowish. Antennae as 
long as entire body; palpi slender, third segment not swollen. Wing 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 77 


with first radial cell a fifth as long as second; petiole of media a fourth 
as long as distal portion. Legs slender, nearly bare, proportions of 
segments of hind leg as in table 1; basitarsi unspined, fifth segment 
with a pair of ventral batonnets. 

Mats: Similar to the female but with the usual sexual differences; 
plumes of antennae yellow. Ninth sternite about four times as broad 
as long, with a very shallow, broad, posterior excavation, the posterior 
membrane spiculate; ninth tergite tapered, much longer than basi- 
styles, with very short, setigerous, apicolateral lobes. Basistyles broad 
at bases, attenuated distad, simple, with an irregular, hyaline mem- 
brane connecting the inner margins about two-thirds way to apices; 
dististyles about as long as basistyles, slightly curved and tapered to a 
slender, pointed tip. Aedeagus with a pair of slender, sclerotized, 
oblique rods with straight apices, the curved bases continuous with 
anterior margins of basistyles. Parameres with bases very stout and 
conical, and with greatly narrowed apices recurved ventrolaterad, the 
bases connected by a narrow, sclerotized band. 

Type.—USNM. 5482, 9, New Jersey, Riverton. 

Material examined.—New Jersey: Riverton, July 3, Johnson, 1 ? 
(type). 

Virginia: Falls Church, July 4, Aug. 6, 1950, Wirth, 4 707, 3 99. 

Georgia: Thomasville, May 15-30, 1949, Palmer, 1 0, 2 99. 


ll. Stilobezzia (Stilobezzia) glauca Macfie, 1939 
FIGuRE 12, h 


Stilobezzia glauca Macfie, Rev. Ent., vol. 10, p. 204, 1939 (o’, Brazil).—Lane, Rev. 
Ent., vol. 18, p. 207, 1947 (92). 

Diagnostic characters —A rather small, bright, pale green species 
with whitish antennae, legs, wings, and halteres; mesonotum exten- 
sively dark, shining brown; abdomen with black band across third 
tergite and lateral spots on fifth. 

Description. —FrEMAueE: Length 1.8 mm., wing 1.7 mm. by 0.6 mm. 
Head yellowish brown, antennae whitish at bases, last five segments 
and palpi pale brownish. Flagellar segments in proportion of 
25:15:15:15:16:17:18:20:40:40:35:35:45. Palpi long, segments in pro- 
portion of 4:7:15:8:12; third segment not swollen, pit absent. 

Thorax pale green; mesonotum with median third of anterior margin 
shining brownish black, forming a broad, V-shaped mark caudad, the 
sides of mesonotum broadly dark to ends of scutellum, and the median 
area between the dark bands yellowish. Vestiture consisting of a few 
long, brown bristles; scutellum with four weak, marginal bristles. 
Postscutellum and a faint pleural spot brown. Legs whitish, hind 
femur and tibia with very faint dark, apical spot. Proportions of 
segments of hind leg as in table 1; basitarsus without spines, fifth seg- 


78 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


ment with a pair of long, dark batonnets at basal fourth; claws very 
slender and unequal, the longer claw as long as fifth segment, the other 
a third as long. 

Wings hyaline, anterior veins yellowish; first radial cell small, a 
seventh as long as second; petiole of media a fifth way to apex of wing. 
Halteres yellowish white. 

Abdomen pale green; two dark spots present on sides of first and 
second tergites; third tergite with broad, black band; fifth tergite 
with a pair of small, lateral, black spots. Spermatheca one, slightly 
oval, the duct not sclerotized. 

Mate: Similar to the female but with the usual sexual differences; 
antennal plumes bright yellow. Ninth sternite broad and not strongly 
sclerotized, with spiculate membrane; ninth tergite rounded with 
membranous, setigerous, apicolateral lobes. Basistyle with distal 
portion slender, base stout, a large unsclerotized lobe about halfway 
along inner margin, connected on dorsal side to margin of ninth tergite; 
dististyle about as long as basistyle, tapered and curved to a slender, 
bluntly pointed tip. Aedeagus with an oblique pair of long, straight, 
slender, sclerotized bars. Parameres with stout basal knobs, a pair 
of inner apodemes joined mesad by a membrane from their anterior 
margins; stems stout, slightly curved, with apex abruptly narrowed, 
pointed, and bent mesad in the form of a short claw. 

Type—tIn collection British Museum (Natural History), o, 
Brazil, Nova Teutonia. 

Material examined.—Virginia: Falls Church, July 4-29, 1950, 
Wirth, 9 oc’, 16 99. 

South Carolina: Greenville, June 20, 1940, H. and M. Townes, 1 oc. 

Louisiana: Baton Rouge, May 1947, Wirth, 2 #0. 

Remarks.—Very similar to S. viridis (Coquillett), 1901, in the gen- 
eral green color with black-banded abdomen, but readily distinguished 
from that species by the dark-banded mesonotum, unbanded legs, 
pale wings, and in the male by the yellowish antennal plumes. One 
female from Falls Church is atypical, with the abdomen entirely dark. 


12. Stilobezzia (Stilobezzia) pallidiventris (Malloch), 1915 


Hartomyia pallidiventris Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 
344, 1915 (9 ; Illinois, Indiana). 
Stilobezzia pallidiventris, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943. 
Description.—FEMALE (from original description): Length 1.0 mm. 
Shining black; head, antennae, palpi, legs, and abdomen yellow; 
abdomen may be darkened; wings hyaline; halteres white. Antennae 
as long as entire body; legs slender; basitarsi longer than segments 
2 to 5 combined; fifth segment without ventral batonnets; claws 
slender, inner half as long as outer. First radial cell a fifth as long as 
second. 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 79 


Type.—tIn collection Illinois Natural History Survey, °, Illinois, 
Urbana. 

Remarks.—Very near S. (S.) diversa (Coquillett), but much smaller, 
the abdomen yellowish rather than green, and fifth tarsal segment of 
female without ventral batonnets. 


13. Stilobezzia (Stilobezzia) pruinosa Wirth, 1952 
FIicureE 12, g 


Stilobezzia pruinosa Wirth, Univ. California Publ. Ent., vol. 9, p. 203, 1952 (7, 

9 ; California). 

Description.—FrmMauE: Length 1.0 mm. Pruinose brown, with 
broad, short, unmarked wings and a marked, pruinose gray pattern 
on the mesonotum; legs with narrow sub-basal pale rings on tibiae; 
halteres white. Proportions of segments of hind leg as in table 1; 
basitarsi and fifth segment without ventral spines. Spermathecae 
two, slightly unequal, slightly ovoid, with a few minute perforations; 
the ducts sclerotized a very short distance. 

Mats: Ninth sternite transverse; ninth tergite long and rounded 
caudad with a subapical pair of low, rounded, setigerous lobes. Basi- 
styles tapered, the inner margins with low hump bearing a few setose 
tubercles; dististyles very stout, with blunt apices. Aedeagus with 
a pair of small, oblique, sclerotized bars. Parameres a pair of highly 
sclerotized, heavy plates; basal apodemes winglike and _ bilobed; 
inner margins of stems nearly straight, slightly expanded on outer 
sides about halfway to apices; apices abruptly narrowed and bent 
ventromesad on about distal fourth. The apices of the parameres 
are somewhat longer in the Florida specimens figured than in the type 
from California. 

Types ——USNM 59945, holotype, o’, allotype, California, San Luis 
Obispo County, Arroyo Seco Ranger Station. 

Material examined.—California: Arroyo Seco Ranger Station, 
San Luis Obispo County, July 1, 1948, Wirth, 3 ao (type, 2 para- 
types), 1 2 (allotype); Pollack Pines, Eldorado County, July 7, 1948, 
Wirth, 1 o&; Snowline Camp, Eldorado County, July 8, 1948, Wirth, 
ire 

Florida: Gainesville, Alachua County, June 2, 1950, Bidlingmayer 
1 o, 1 9; Grayton Beach, Walton County, May 10, 1949, Butler, 1 9 


14. Stilobezzia (Stilobezzia) punctipes, new species 
FicureE 11, d 


Diagonostic characters.—A large, yellowish species with brown 
thorax and brown-punctate legs; abdomen with narrow segmental 
dark bands; wings with two small black spots. 


80 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Description.—FEMALE: Length 2.4 mm., wing 1.9 mm. by 0.9 mm. 
Head pruinose brown; antennae yellow, apices of segments 3 to 10 
and all of last five segments brown; palpi black. Flagellar segments 
in proportion of 28:18:18:18:18:18:20:20:38:38:40:40:55, last 
segment with very slender pointed tip. Papal segments in proportion 
of 5:10:16:8:20, third segment slightly swollen with a small, round, 
subapical pit. 

Thorax pruinose brown, margins of mesonotum, broad center of 
scutellum, and upper half of pleura yellow. Mesonotum with irregular, 
silvery pruinose patches, a few long, brown bristles; scutellum with 
four long, marginal bristles. Coxae and trochanters dark brown, 
rest of legs yellow; fore femur with broad sub-basal, midfemur with 
narrow basal, and hind femur with narrow apical brown bands; all 
tibiae narrowly brown at apices; rows of small, round, brown puncta- 
tions at bases of hairs on all femora and tibiae, these hairs long and 
dark on hind tibia. Proportions of segments of hind leg as in table 
1; basitarsus with three black ventral spines on forelegs, two on 
midlegs and one at base on hind legs; fifth tarsal segment without 
batonnets; claws slender and unequal, the longer claw as long as 
fifth segment, the other two-thirds as long. 

Wing hyaline, anterior veins yellowish; a brown spot over r-m 
crossvein and a very small one over vein R:,3; macrotrichiae absent. 
First radial cell large, one-fourth as long as second, latter to 0.8 wing 
length; petiole of media as long as r—m crossvein. Halteres brown. 

Abdomen dull, pruinose yellowish, tergites with irregular brown 
bands in middle of segments, pleura dark. Spermathecae three, the 
two larger subequal and oval with short, sclerotized ducts, the third 
about two-thirds as large and subspherical, without sclerotized duct. 

Mate: Similar to the female but with the usual sexual differences; 
plumes of antennae golden brown. Ninth sternite a very narrow band, 
the posterior membrane spiculate; ninth tergite rounded, with small, 
submedian, setigerous lobes at apex. Basistyle simple, twice as long 
as broad; dististyle as long as basistyle and rather stout to apex, 
slightly bent on distal third. Aedeagus with an oblique pair of heavy 
twice-sinuate sclerites with apices snoutlike and crossing each other 
mesad. Parameres very slender, the lateral apodemes evenly curved 
and slender, stems nearly straight with bases slightly enlarged, 
gradually tapered to sharp, hooked tips abruptly bent ventrocephalad 
at two-thirds total length. 

Types—USNM 60966, holotype, ?, Crystal River, Citrus County, 
Fla., June 7, 1949, Hudson, light trap; allotype, same data except 
date, Oct. 6, 1949. Paratypes: Florida: 11 @o, 23 99, same data 
as type except dates, May-October 1949-50; 167, 3 99, Leesburg, Lake 
County, Aug. 7, 1949, Braddock; México: .4 99, Ciudad Monte, 
Tamaulipas, Nov. 22, 1943, Brookman. 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 81 


Most closely related to the Neotropical species S. (S.) punctulata 
Lane, 1947, and S. (S.) kiefferi Lane, 1947, which it resembles in its 
spotted wings and punctate legs. Both related species, however, 
have the abdomen reddish, the former species differing in having 
three dark wing spots, the third over tip of second radial cell, while 
the second species has the first radial cell half as long as the second, 
yellowish coxae, whitish halteres, and the fore femur and midfemur 
lack the basal dark bands. 


15. Stilobezzia (Stilobezzia) rabelloi Lane, 1947 
FIGureE 11, a 


Stilobezzia rabellot Lane, Rev. Ent., vol. 18, p. 203, 1947 (o, 9 ; Brazil). 


Diagnostic characters——A large, brown species with irregular, 
silvery, pruinose markings; wing brownish, infuscated with three 
darker and one lighter spots on anterior margin; body hairs dense and 
strong; fifth tarsal segment with 3 or 4 pairs of ventral batonnets. 

Description —Frmate. Length 3.0 mm., wing 2.2 mm. by 0.8 mm. 
Head pruinose brown, narrow bases of flagellar segments yellow, 
palpi black. Antenna with flagellar segments in proportion of 
22:15:15:15:18:20:24:24:40:42:50:?:? (last two segments broken, 
not measured). Palpal segments in proportion of 8:15:25:15:20, 
third segment slightly swollen, densely set with strong hairs, a small, 
round pit near apex. 

Mesonotum with strong, conical, anterior spine, the sides sloping, 
flattened only in prescutellar area; color dark brown with irregular 
silvery pruinose markings, usually three sublateral pairs of small, 
rounded, polished spots on disc; humeral corners yellowish; a few long, 
brown hairs in row. Scutellum brown, yellowish in middle, with 
4 or 5 long, brown, marginal bristles. Postscutellum and pleura dark, 
pollinose brown. Legs including coxae dark brown; fore femur and 
midfemur with narrow sub-basal and subapical and broad median 
yellowish bands; hind femur with narrow, light band at apex; fore 
tibia and midtibia with narrow sub-basal and broad median yellowish 
bands; first four tarsal segments yellowish. Proportions of segments 
of hind leg as in table 1; leg hairs strong and spinose, especially long 
on hind tibia; basitarsus with strong black basal and apical spines, a 
third spine in middle on forelegs; fifth segment with 3 or 4 pairs of 
ventral batonnets; claws very slender and unequal, the longer claw 
as long as fifth segment, the other a third as long. 

Wing brownish hyaline, extensively infuscated along the veins; 
three dark brown spots, first over r-m crossvein, second over vein 
R243, and the third over distal third of second radial cell and extending 
halfway through cell R,; at same level; a light area just beyond tip 


82 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


of costa as large as third brown spot, the apex of the wing beyond 
lightly infuscated. First radial cell nearly half as long as second, the 
latter ending at about 0.7 wing length; petiole of media about as long 
as first radial cell. Halteres brown except at extreme bases of stems. 

Abdomen dark brown, tergites with narrow, apical, segmental, 
silvery pruinose bands; vestiture very dense, consisting of long, stout, 
brown hairs. Spermathecae two, ovoid, subequal, with ducts sclero- 
tized a short distance, and a rudimentary third present. 

Maus: Similar to the female but with the usual sexual differences; 
plumes of antennae brown. Ninth sternite about three times as 
broad as long, with shallow, mesal, posterior excavation, and the 
membrane bare; ninth tergite short and rounded with very short, 
setigerous, apicolateral lobes. Basistyles simple, very long, slender 
and tapered to tips, about three times as long as broad; dististyles 
about 0.8 as long as basistyles, slightly incurved, slender, with pointed 
apices. Aedeagus with an oblique pair of double-sinuate sclerites 
with mesal tips snoutlike. Parameres strongly sclerotized with 
broad, winglike, lateral apodemes at bases, a lateral pair of L-shaped 
or boomerang-shaped lateral plates with blunt apices directed latero- 
caudad, and a submedian pair of nearly straight processes of same 
thickness, but about twice as long, with pointed apices. 

Type.—In collection University of Sao Paulo, Brazil, @, Brazil, 
Rio de Janeiro, Estrada Rio-Séo Paulo, km. 47. 

Material examined.—Georgia: Thomasville, May 15-30, 1949, 
Palmer, 1 9. 

Florida: Leesburg, Lake County, Aug. 7, 1949, Braddock, 2 99; 
Panama City Beach, May 6, 1949, McElvey, 3 <7, 1 2; Santa Rosa 
Island, Escambia County, June 7, 1949, Butler, 1 #7, 1 9. 

Louisiana: New Orleans, May 27, 1947, Van Beeck, 6 co. 

Honduras: La Ceiba, Aug. 6, 1916, Dyer, 1 &. 

Surinam: Moengo, Nov. 10, 1947, Stage, 1 9. 

Remarks.—Closely related to S. (S.) beckae, new species, which it 
resembles closely in the brown mesonotum with silvery pruinosity and 
strong anterior spinose tubercle and brownish infuscated and spotted 
wings, but beckae differs in lacking the third dark spot at apex of first 
radial cell and the ventral batonnets on fifth tarsal segment, and the 
body hairs are not nearly so well developed. 


16. Stilobezzia (Stilobezzia) sybleae, new species 
FIGuRE 12, c 


Diagnostic characters.—A small, shining, brown species with dusky, 
yellow legs; wing hyaline with a prominent beadlike swelling of vein 
Ra;s near apex of second radial cell; hind tarsi with fifth segment and 
claws greatly reduced. 


HELEID MIDGES, GENUS STILOBEZZIA—-WIRTH 83 


Description.—FEMALE: Length 1.5 mm., wing 1.3 mm. by 0.5 mm. 
Head shining brown, clypeus and antennal pedicel yellowish, antennal 
flagellum and palpi dusky. Thorax polished brown, mesonotum with 
sparse black hairs in rows; scutellum with four marginal hairs. Pleura, 
coxae, midfemur, hind femur, and tibiae dusky, yellowish brown; all 
of forelegs, midtarsus, and hind tarsus lighter yellowish. Proportions 
of segments of hind leg as in table 1; basitarsi unspined; fifth segment 
with a pair of black ventral batonnets near base; claws slender and 
unequal, normal on forelegs and midlegs with outer claw as long as 
fifth segment, the inner a third as long, on hind legs with fifth seement 
half as long and claws only a fourth as long as on forelegs and midlegs. 

Wings grayish hyaline, without macrotrichiae, anterior veins brown- 
ish yellow. First radial cell a fourth as long as second, latter to five- 
eighths wing length; vein Ry; on distal half of second radial cell with 
an oval beadlike swelling; crossvein r-m short, petiole of media about 
three times as long as crossvein. Halteres brown. Abdomen shining 
brown, very convex dorsally and somewhat petiolate at base. 

Mate: Ninth sternite a slender anterior band, the posterior mem- 
brane spiculate; ninth tergite rounded, the apicolateral lobes reduced. 
Basistyle broad, with a large triangular, flattened, sclerotized lobe 
about midway of mesal margin; dististyle nearly as long as basistyle, 
incurved, gradually tapered to slender apex. Aedeagus with a pair of 
slightly oblique, long, very slender, sclerotized bars. Parameres with 
basal knoblike apodemes fused with anteromesal margins of inner 
lobes of basistyles, with slender, long, curved, sicklelike posterior 
blades. 

Types ——USNM 60967, holotype, 9, and allotype, Falls Church, 
Virginia, July 4, 1950, Wirth. 

Remarks.—This species is very similar to S. (S.) antennalis (Coquil- 
lett), but the highly modified posterior branch of the radius and hind 
fifth tarsal segment and claws are quite distinctive. The West 
African species S. limnophila Ingram and Macfie, 1922, is very similar 
in coloration and in the structure of the male genitalia, but since the 
female is unknown, the wing and tarsal features cannot be compared 
with those of sybleae. Parabezzia poikiloptera Ingram and Macfie, 
1922, from West Africa, which is a Stilobezzia of the subgenus Hukrai- 
ohelea, has a remarkable thickening of the radius almost exactly like 
that of sybleae, indicating that this character is not just an aberration. 
This species is named for my wife, Syble Austin Wirth. 


17. Stilobezzia (Stilobezzia) thomsenae, new species 
FIGURE 12, a 


Diagnostic characters.—Very closely related to, and almost indis- 
tinguishable from, bulla Thomsen, except for the very characteristic 
male genitalia. 


84 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Description.—FEMALE: Length 1.0 mm.; wing 0.8 mm. by 0.4 mm. 
Apparently somewhat darker brown than bulla, the mesonotum and 
scutellum concolorous; femora brownish. Otherwise as in bulla. 

Mate: Genitalia broader than long, parts very broad and stout. 
Ninth sternite a narrow, anterior band, posterior membrane spiculate; 
ninth tergite rounded, with a submedian pair of fingerlike setose lobes 
about twice as long as broad. Basistyles simple, about twice as long as 
broad, set far apart, no inner lobes; dististyles short and stout, two- 
thirds as long as basistyle, with abruptly narrowed, pointed, clawlike 
apices. Aedeagus with anterior arms vestigial, the dorsal median 
sclerite a crescent-shaped, transverse bar connecting distoventral 
angles of basistyles. Parameres stout, with broad, lateral, winglike, 
basal apodemes, stems slightly knobbed at bases, stout and slightly 
out-curved toward apices, which are slightly flattened, expanded, 
rounded, and bent ventrad. 

Type-—USNM 60968, holotype, @, Everglades City, Collier 
County, Fla., Feb. 7, 1950, Davidson, light trap. 

Remarks.—One female from Crystal River, Citrus County, Fla., 
Sept. 18, 1950, Hudson, is provisionally referred to S. (S.) thomsenae, 
since it resembles the male in coloration, although it may prove to be 
a dark specimen of S. (S.) bulla. This species is named in honor of 
Dr. Lillian Thomsen, who has contributed much to the study of 
North American Heleidae. 


18. Stilobezzia (Stilobezzia) viridis (Coquillett), 1901 
Figure 12, d 

Ceratopogon viridis Coquillett, Proc. U. 8. Nat. Mus., vol. 23, p. 607, 1901 (9, 

New Jersey). 
Ceratolophus viridis, Kieffer, Genera insectorum, fase. 42, p. 61, 1906. 
Johannseniella viridis, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 

227, 1914. 
Hartomyia viridis, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 342, 

1915. 
Stilobezzia viridis, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1948. 

Description.—FEMAuE: Length 2.0 mm., wing 1.9 mm. by 0.6 mm. 
A bright, pale-green species; antennae and legs yellow, apex of antenna 
dark; distal fourth of hind femur and narrow apex of hind tibia black; 
abdomen with a black band across distal half of third tergite and a 
pair of large black spots on fifth tergite. Wings brownish hyaline, 
anterior veins infuscated. Proportions of segments of hind leg as in 
table 1; basitarsi without spines; fifth segment with a pair of long, 
black batonnets at basal fourth; claws slender and very unequal, the 
longer as long as fifth segment, the other about half as long. 
Mate: As in the female but with the usual sexual differences, 


HELEID MIDGES, GENUS STILOBEZZIA—WIRTH 85 


plumes of antennae dark. Genitalia very large and bulbous. Ninth 
sternite reduced, the posterior membrane spiculate; ninth tergite 
large, convex, apex rounded with low, rounded, setigerous, apicolateral 
lobes. Basistyle short and very broad, nearly as broad as long; 
dististyle about 1.2 times as long as basistyle, extreme base bulbous, 
distal portion very slender, irregularly bent inward, with blunt apex 
slightly bent outward. Aedeagus with a pair of slender, nearly 
straight, barlike sclerites. Parameres with reduced basal scleroti- 
zation, stems slender and slightly crooked, apices greatly expanded in 
a rounded lobe. 

Type-—USNM 5483, 9, New Jersey, Riverton. 

Material examined—New Jersey: Riverton, June 16, Johnson 
(type). 

Maryland: Cabin John, May 26, 1943, Cortes and Townes, 2 07; 
Glen Echo, June 30, 1929, Bridwell, 1 o. 

Florida: Leesburg, Lake County, Aug. 7, 1949, Braddock, 1 9. 

Texas: Kirbyville, May 20, 1908, Tucker, 1c. 

Remarks.—S. glauca Macfie from Brazil and the southeastern 
United States is closely related to viridis, but has the antennae and 
wings infuscated, the mesonotum dark, and lacks the apical black 
markings on the hind femur. 


References 


GoETGHEBUER, M. 
1934. Heleidae (Ceratopogonidae). Jn Lindner, Die Fliegen der Palal- 
arktischen Region, Lief. 78, pp. 49-94. 
JOHANNSEN, O. A. 
1934. New species of North American Ceratopogonidae and Chironomidae. 
Journ. New York Ent. Soc., vol. 42, pp. 343-353. 
1943a. Two new species of American Ceratopogonidae (Diptera). Ann. 
Ent. Soc. Amer., vol. 36, pp. 761-762. 
1943b. A generic synopsis of the Ceratopogonidae (Heleidae) of the Americas, 
a bibliography, and a list of the North American species. Ann. 
Ent. Soc. Amer., vol. 36, pp. 763-791. 
1946. Some new species of Nemocerous Diptera from Guam. Bull. Bishop 
Mus., No. 189, pp. 189-193. 
LANE, J. 
1947. Espécies Brasileiras de Stilobezzia (Dipt., Ceratopogonidae) e Zygone- 
ura stonei nov. nom. (Dipt., Mycetophilidae). Rev. Ent., vol. 18, 
pp. 197-214, 9 figs. 
Macrig, J. W.S. 
1935. Ceratopogonidae (Dipt.) from the river Amazon. Stylops, vol. 4, 
pp. 49-56, 2 figs. 
MEILLON, B. DE 
1938. Notes on African Ceratopogonidae (Diptera). Proc. Ent. Soc. 
London, ser. B, vol 7, pp. 266-270. 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


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SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1953 No. 3317 


BEETLES OF THE OEDEMERID GENUS VASACES 
CHAMPION 


By Ross H. Arnett, Jr.’ 


Champion (1889) described the genus Vasaces for three species from 
Central America. One species, Vasaces aeneipennis, was described 
from material from México, and the other two, Vasaces sordidus and 
Vasaces costatus, were described from Guatemala. 

Since the completion of my revision of the Nearctic Oedemeridae 
(Amer. Midl. Nat., vol. 45, pp. 257-391, 1951), I have seen several 
specimens of Vasaces collected by J. N. Knull in Arizona and Texas. 
This genus has not previously been recorded from North America, 
north of México. In addition, a few specimens from Central America, 
in the collections of the U. S. National Museum and the Museum of 
Comparative Zoology, have been available to me for study. 

The present paper, based on all available material, with the excep- 
tion of the Champion type specimens, adds four new species to the 
genus. <A key to the species is included, and the position of the genus 
in relation to other oedemerid genera is discussed. 

I wish to thank Mr. J. N. Knull, Ohio State University, and Dr. 
P. J. Darlington, Museum of Comparative Zoology, for the loan of 
material used in this study. 


1Bureau of Entomology and Plant Quarantine, U. S. Department of Agriculture. 


232714—53 87 


88 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Genus Vasaces Champion, 1889 


Vasaces Champion, Biologia Centrali-Americana, Coleoptera, vol. 4, pt. 2., p. 111, 
1889 [no included species] and p. 128, pl. 6, figs. 7 and 7, a-c, 1890 [first in- 
cluded species].— Arnett, Journ. Washington Acad. Sci., vol. 40, p. 225, 1950. 
(Genotype, Vasaces aeneipennis Champion, 1890; designated by Arnett, 
1950.) 

This genus belongs in the subfamily Oedemerinae as treated in my 
paper on the Nearctic species. The apical segment of the maxillary 
palpus is triangular or cultriform and the fore tibia has two apical 
spurs. It is tentatively placed in the tribe Asclerini, as I have pre- 
viously defined it. However, studies of certain other genera have 
led me to revise somewhat my views on the oedemerid tribes, and 
when more groups have been restudied some rearrangement may be 
necessary. Vasaces Champion will key to couplet 7 of my key to the 
genera of Asclerini. The slightly emarginate eyes separate it from 
Heliocis Arnett, and the triangular or cultriform apical segment of the 
maxillary palpus will separate it from the genus Sisenes Champion. 
In Sisenes Champion the apical segment of the maxillary palpus is 
small, hardly triangular. In addition, many Sisenes species resemble 
lycids and have the antennal segments somewhat flattened, a condi- 
tion which is not found in Vasaces. In the key to the male genitalia, 
this genus runs to couplet 3, where it may be separated from the 
genera Humecomera Arnett and Sisenes on the basis of the structure 
of the ninth abdominal sternite. In Vasaces the apical portion of the 
ninth sternite of both Eumecomera and Sisenes consists of two separate 
rods united only by thin membrane. The ninth sternite of Vasaces 
resembles somewhat that of Heliocis except that the base is two 
distinct rods instead of the Y-shaped piece found in Helvocis. 

It appears that the genus Sisenes is a close relative of Vasaces, 
both on characters of external morphology and those of the male 
genitalia. However, in the studied forms, the two genera can be 
adequately separated using the characters mentioned above. In 
gross appearance, the two genera are very distinct. The species of 
Vasaces are for the most part dark, somewhat metallic, elongate 
beetles, quite different from the brilliantly colored, lycidlike species 
of Sisenes. 

Generic diagnosis—Head moderate or elongate, usually constricted 
behind the eyes. Eyes somewhat emarginate, large. Antenna 
slender, 11-segmented, inserted between the eye and the base of the 
mandible, distinctly separated from the margin of the eye and the 
base of the mandible. Second segment of the antenna large, over 
one-half the length of the fourth and sometimes nearly the length of 


OEDEMERID BEETLES—ARNETT 89 


the third. Both mandibles with the apex of each bifid. Apical 
segment of the maxillary palpus large, triangular or cultriform, widest 
at the base or near the center. Thorax slightly longer than broad, 
usually only slightly expanded apically and only slightly constricted 
at the base. Legs slender, claws simple; all tarsi with only the 
penultimate segment of each spongy beneath. Elytra distinctly 
costate, sometimes markedly so. 

Mae GeniTALia: Median lobe short, base wider, without proc- 
esses, tapering towards apex, without apical enlargements. Paramere 
a troughlike plate, wide basally, without basal processes, narrowed 
toward apex; apex V-shaped, emarginate; tegminite large, spatulate. 
Ninth abdominal sternite hood-shaped, with two basal tails; apex 
expanded, with basally projecting lateral flanges. Ninth tergite 
two parallel rods, apically expanded and united by membrane. 
Eighth abdominal sternite and tergite each bilobed, the sternite larger. 
Setae present on the apical lobes of the eighth abdominal segment. 

Biology —Unknown. 

Distribution Known only from Central America and the extreme 
southwestern part of the United States. 


Key to the species of Vasaces 


1. Apical segment of the maxillary palpus cultriform (fig. 13,g), its outer edge 
SIMIC era ere LE SAE. Oy RPE OT RE SER ote ee ee SR RS eS 2 
Apical segment of the maxillary palpus triangular (fig. 13,h)--.------------ 4 

2. Costae, suture, and margins of elytra fusco-piceous, intervals pale; antennae 
uniformly dark except for small pale areas at the joints; legs dark, 


Rep AICr AG JOUS. 222 A 8 ee linearis, new species 
Vinee OOM IOrMCOlOTAuiOUe | = o..e2 oo same = oe ae Se ee 3 

pe tavernainitormily brassyweszt eswes 8- ical il dudes costatus Champion 
BN rab etn eur Ke eres SR en ae knulli, new species 

4. Pale testaceous; elytra with costae and suture each with an indistinct, inter- 
rupted row of fine, oblong streaks on either side___-_-- sordidus Champion 
Mee eeeics, OGUerwitt MATKCOL. 20 Sh to ee oe eee eee 5 

5. Pronotum orange with black maculations, head short__maculatus, new species 
napataniadark, head elongate: sos) Soi th. oot ee om al oa 6 

6. Pronotum with sides nearly parallel____._.-._---------- elongatus, new species 
Pronotum distinctly sinuate laterally____..._-___-.- aenipennis, Champion 


Vasaces linearis, new species 
FicurE 18, c, e-g, j-l, m, n 
Diagnostic characters —The light brassy color, with the dark brown, 
prominent elytral costae, readily differentiates this species from the 


others. It is most closely related to V. costatus since the shape of 
the apical segment of the maxillary palpus is similar. 


90 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Description.—(Based on male holotype.) Antenna long, nearly as 
long as the body, second segment subequal to the third. Apical 
segment of the maxillary palpus large, distinctly cultriform, outer 
margin arcuate. Vertex of head transversely elevated between the 
eyes, and with a slight median groove from the base of the antenna 
to the summit of the vertex. Pronotum somewhat longer than broad, 
widest near anterior margin, slightly narrower at base, surface irregu- 
lar, laterally with slight elevations, median line somewhat impressed, 
and with a slight V-shaped elevation on anterior half. Elytron long 
and narrow, with four prominent costae and the suture and margin 
prominently elevated, and a short, basal, spurious costa uniting with 
the inner margin. 

Entire insect clothed with fine, short, sparse, whitish hairs. Color: 
in general pale brassy; head with a darker area on front; mouthparts, 
except for maxillary palpi, paler. Antennae and maxillary palpi dark. 
Thorax dark except for irregular paler areas near base and apex. 
Elytra uniformly pale brassy except for dark brown costae, suture, 
and margins. Ventral surface and legs dark. Surface irregularly 
rugose-punctate. Length: 8 mm. 

Mate GeniTatia: Described in the generic diagnosis. The seventh 
sternite has a slight angular emargination which is probably a second- 
ary sexual character. 

Type.—Holotype, male, Ohio State Univ. coll., Arizona, Huachuca 
Mountains, Cochise County, June 9, 1935, J. N. Knull. 

Distribution.—Known only from this single male type specimen 
from Arizona. 

Vasaces costatus Champion, 1890 
Vasaces costatus Champion, Biologia Centrali-Americana, Coleoptera, vol. 4, 
pt. 2, p. 129, pl. 6, fig. 8, 1890. 

This species is also known only from one male, the type, from 
Guatemala, at Cerro Zunil (4,000 feet), collected by Champion, and 
is in the British Museum (Natural History). It is much smaller 
than the genotype, but a little larger (9.5 mm.) than the preceding 
species. The elytra are uniformly brassy in color, lacking the dark 
brown costal, sutural, and marginal markings. The seventh sternite 
of the male is deeply emarginate. 


Vasaces knulli, new species 
FicureE 13, b 
_ Diagnostic characters—The small size, dark color, and lack of 
brassy tints serve to separate this species from the other two having the 


apical segment of the maxillary palpus cultriform. It is most closely 
allied to V. costatus. 


OEDEMERID BEETLES—ARNETT 91 


Ficure 13.—Adults and male genitalia of the genus Vasaces: a, Vasaces maculatus, new 
species, 2 ;b, V. knulli, new species, 2 ; c, V. linearis, new species, o'; d, V. elongatus, new 
species, 9., ¢, f, co genitalia, V. linearis, new species: e¢, paramere; f, median. lobe; g, A, 
maxillary palpi: g, /. linearis, new species, o'; h, V. elongatus, new species, 9 ; 1, V. 
aeneipennis Champion, 9; j—-n, V. linearis, new species, 7; J, ninth tergite; &, ninth ster- 
nite; /, tegminite; m, eighth tergite; , eighth sternite. 


92 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Description.—(Based on female holotype.) Antennae short, not 
extending over two-thirds the length of the elytra, second and third 
segments equal in length, each about three-fourths the length of the 
fourth segment. Apical segment of the maxillary palpus large, 
cultriform, outer edge slightly arcuate. Head shorter than that of the 
other species; front with a slight depression near the bases of the 
antennae on the inner side; a moderately deeply impressed median 
line on the front. Pronotum with surface very irregular and with a 
moderately prominent V-shaped ridge on the anterior half and 
an interrupted lateral ridge on each side; longer than broad, widest 
anteriorly just beyond the middle, constricted behind. Elytra nar- 
row, long; costae present but obscure. Surface covered with fine, 
short, sparse, whitish hairs. Color: dark, mouthparts, underside 
of head, neck, coxae, bases of femora, tarsi at joints, and small area at 
joints of the segments of the antennae paler. Surface finely rugose- 
punctate. Length: 7 mm. 

Type.—Holotype, female, Ohio State Univ. coll., Arizona, Huachuca 
Mountains, Cochise County, June 9, 1935, J. N. Knull. 

Distribution.—This species is known only from the type, which is 
from Arizona. 

Vasaces sordidus, Champion, 1890 
Vasaces sordidus Champion, Biologia Centrali-Americana, Coleoptera, vol. 4, pt. 
2, p. 129, 1890. 

This species is known from one female only, the type, from Capetillo, 
Guatemala, collected also by Champion, which is in the British Mu- 
seum (Natural History). It is smaller than the preceding species 
(8.5 mm.); the apical segment of the maxillary palpus is smaller, and 
triangular instead of cultriform. The color is pale testaceous. The 
costae and suture are conspicuous, and each has an indistinct, inter- 
rupted row of fine oblong streaks on either side. 


Vasaces maculatus, new species 
FIGuRE 13, a 


Diagnostic characters —The dark orange pronotum with a median and 
two lateral spots is unique among the species of this genus. It is the 
largest known species in the genus and is most closely related to V. 
sordidus but has the orange markings which are lacking in that species. 

Description.—(Based on female holotype.) Antennae short, reach- 
ing to about two-thirds the length of the elytra, second segment short, 
about two-thirds as long as the third. Apical segment of the maxillary 
palpus large, elongate triangular, widest near basal third, outer mar- 
gin straight. Front of head irregular, with a circular depression 
extending from the inner side of the base of the antenna basad about 
half the distance to the pronotum. Pronotum quadrate, widest 


OEDEMERID BEETLES—ARNETT 93 


anteriorly, only slightly constricted posteriorly, surface irregular but 
without prominent ridges or depressions except for a slight depression 
on either side near the anterior lateral angle. Elytra long, narrower 
apically, about twice as broad as thorax; costae present, flat, not promi- 
nent. 

Entire insect clothed with very fine, short, sparse, golden hairs. 
Surface finely rugose-punctate. Head with punctures larger, more 
distinct, with a stouter recumbent hair arising near each. Color: 
Head orange, with a large dark spot on front; antennae and palpi dark; 
thorax orange except for a large central dark spot which connects with 
two large lateral spots; legs with coxae and bases of femora pale, re- 
mainder dark; abdomen and elytra uniformly dark. Length: 10mm.; 
of paratype, 11 mm. 

Types.—Holotype, female, USNM (61112, Arizona, Chiricahua 
Mountains, June 8. Hubbard and Schwarz. Paratype, female, same 
data as type except date, June 11. 

The paratype agrees with the type except that the black markings 
of the thorax are less extensive. 

Distribution—Known only from the two type specimens from 
Arizona. 

Vasaces elongatus, new species 


Ficure 13, d, h 


Diagnostic characters.—The uniform dark color, elongate head and 
thorax, and parallel sides of the pronotum are characteristic of this 
species. Itis very different in general habitus from the other members 
of this genus, but is probably most closely related to V. aeneipennis. 

Description.—Head elongate, over twice as long as wide at bases 
of antennae, hardly constricted behind the eyes. Antennae reaching 
beyond apical third of elytra, apical segment of the maxillary palpus 
triangular, widest at middle, outer edge straight. Front of head 
smooth, without elevations or depressions. Pronotum longer than 
broad (7:4), surface irregular but without distinct ridges or depressions, 
sides only slightly sinuate. Elytra broad, twice the width of the 
thorax, tapering behind. Legs long and slender, costae distinct, flat. 

Entire insect clothed with short, fine, white hairs, those of the head 
and thorax sparser, but longer and coarser than those of the rest of 
the body. Surface of head smooth and shiny with large distinct 
punctures which are sparser on the vertex; thorax coarsely rugose; 
remainder of body finely rugose. Color: dark fuscous; neck, under 
surface of head, mouthparts, and coxae paler. Length: 8 mm. 

Type.—Holotype, female, Ohio State Univ. coll., Texas, Chisos 
Mountains, Brewster County, June 9, 1939, D. J. and J. N. Knull. 

Distribution.—Known only from the female type. 


94 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Vasaces aeneipennis Champion 
FIGuReE 18, 7 


Vasaces aenipennis Champion, Biologia Centrali-Americana, Coleoptera, vol. 4, 
pt. 2, p. 128, 1890. 

This is apparently the least rare of the species in this genus and 
is one of the largest of the known species. The triangular apical 
segment of the maxillary palpus, the elongate head, and the uniform 
brassy color distinguish it from the other species. 

Champion described this species from two specimens (a male and 
a female) from Totosinapan, México, in the Sallé collection (location 
unknown). The U.S. National Museum collection contains two fe- 
males from Guatemala, and the Museum of Comparative Zoology has 
one female from Honduras. A detailed description of one female 
from Guatemala follows: 

Description.—(Based on female, from Guatemala). Head elongate, 
about twice as long as wide at bases of antennae. Antennae long, 
reaching nearly to apical third of the elytra, second segment about 
two-thirds the length of the third. Apical segment of the maxillary 
palpus triangular, widest at middle, outer edge straight. Dorsal 
surface of head irregular, but without distinct ridges or depressions. 
Pronotum nearly quadrate, widest anteriorly, sides distinctly sinuate, 
surface irregular. Elytra long and narrow, tapering at apex; costae 
distinct, flat, both edges of costae outlined by a very fine, irregular, 
impressed line, somewhat darker in color, giving in gross appearance 
the impression that the costae are double. 

Surface clothed with extremely fine, very short, sparse, golden 
hairs which are somewhat larger on the thorax and dorsal surface of 
the head. Surface rugose, that of the thorax and head indistinctly 
rugose-punctate. Color: uniformly brassy, dorsal surface of the head 
and ventral surface of the thorax and abdomen darker. Length: 
15 mm. 

Variation: The surface of the head and thorax varies from rugose- 
punctate to shiny smooth with sparse punctures. The length varies 
from 10 to 16 mm. The color is darker in some specimens than in 
others. 

Distribution—Known only from the types from México and the 
above-mentioned specimens from Guatemala and Honduras (Rosario 
San Juancito). 

Material examined.—Female, USNM coll., Guatemala, Santa 
Maria, W. Schaus. 


U. S. GOVERNMENT PRINTING OFFICE: 1953 


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Vol. 103 Washington : 1953 No. 3318 


SCARABAEID BEETLES OF THE GENUS BRADYCINETULUS 
AND CLOSELY RELATED GENERA IN THE UNITED STATES 


By O. L. Cartwricut 


The group of beetles reviewed in this paper have long been known 
under the name Bolboceras. The species found in the United States 
were last revised by Charles Schaeffer, 1906 (Trans. Amer. Ent. Soc., 
vol. 32, pp. 249-254). At that time five species were placed in Horn’s 
genus Bradycinetus, later incorrectly synonymized under Bolboceras 
(see synonymy under Bradycinetulus), and three were assigned to 
Bolboceras. Of these three, Scarabaeus lazarus Fabricius had been 
placed in Hucanthus by Westwood, 1848 (Proc. Linn. Soc. London, 
vol. 1, p. 387), and Bradycinetus hornw Rivers and B. minor Linell 
were later removed to Kolbeus by Chapin (see synonymy under 
Bolbelasmus). 

With the exception of the species Hucanthus and Bolbocerosoma, here 
considered correctly placed in recent usage, all of the United States 
species listed by Schaeffer have been reexamined in this study and 
reassigned to genera as required by the literature on the group. Also 
included in this paper are the one species described since Schaeffer’s 
time, Bolboceras angulus Robinson; a Mexican species discovered in the 
United States; another which may eventually be found inside our 
borders; and four new United States species. Two new genera are 
proposed. A key, based in part on that by Boucomont, is presented 
for the genera of the Bolboceratini found in the United States. The 
genera Kolbeus, Bolbelasmus, Bradycinetulus, and the two new genera 
Bolbocerastes and Bolborhombus are discussed, synonymy listed, and a 
key to the species given. Pertinent references, the type locality and 
location of the type, and notes on distribution follow for each species. 
Distribution is also shown on the maps, figures 15 and 16. The genus 

232990—53——1 95 


96 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Bolbocerosoma, revised by Dawson and McColloch, 1924 (Canadian 
Ent., vol. 56, pp. 9-15), is not included, nor is the true Bolboceras, 
which was revised under the name QOdontaeus by J. B. Wallis, 1928 
(Canadian Ent., vol. 60, pp. 119-128, 151-156, 168-176). 

The number of specimens available for study was greatly increased by 
material borrowed from museum and private collections. I am grate- 
ful for the opportunity to study these added specimens and thank the 
following who loaned museum specimens: R. H. Beamer, Snow 
Museum, University of Kansas (KAN); E. C. Becker, Canadian 
Department of Agriculture (CDA); M. A. Cazier, American Museum 
of Natural History (AMNH); P. J. Darlington, Museum of Compara- 
tive Zoology (MCZ); Henry Dietrich, Cornell University (CU); T. H. 
Hubbell, University of Michigan (MICH); P. D. Hurd, University 
of California (CAL); J. N. Knull, Ohio State University (OSU) ; Hugh 
B. Leech, California Academy of Sciences (CAS); L. M. Martin, Los 
Angeles County Museum (LAC); H. J. Reinhard, A. & M. College of 
Texas (TEX); V. D. Roth, Oregon State College (ORE); M. W. 
Sanderson, Illinois Natural History Survey (INHS); and G. E. 
Wallace, Carnegie Museum, Pittsburgh (CARN). My thanks are 
extended also to those who loaned specimens from their private collec- 
tions: R. H. Arnett, D. K. Duncan, C. A. Frost, P. C. Grassman, 
Henry Howden, A. T. McClay, G. H. Nelson, F. H. Parker, Mark 
Robinson, E. C. VanDyke, and F. G. Werner. I am especially grate- 
ful to Henry Howden who examined my types, compared specimens 
with them and later furnished those records marked with an asterisk. 
The few records of the eastern Bradycinetulus ferrugineus (Palisot de 
Beauvois) taken from the literature are marked with a double asterisk. 
Florida records from the literature are from Blatchley (Florida Ent., 
vol. 12, p. 29, 1925), Alabama records from Léding (Geol. Surv. 
Alabama Monogr. 11, p. 101, 1945), and North Carolina records from 
Brimley (The insects of North Carolina, p. 201, 1938). 

All other distribution records were copied from labels attached 
to specimens I examined. 


Key to the genera of Bolboceratini 


1: Eye entirely ‘divided by ocular canthus. . .-- +122. 5.2 2 2 eee 2 
Eye not completely divided by ocular canthus___---.------------------ 3 

2. Middle coxae narrowly separated; the intercoxal process with a vertical tooth- 
like elevation in front of the coxae; species usually bicolored__Bolbocerosoma 

The intercoxal process without toothlike elevation; species not bicolored; 
males usually with long cephalic horn—-__-_-----2 2-2-2 eee Bolboceras ! 

3. Middle coxae subcontiguous; metasternum linear between the coxae_ -__-- + 
Middle coxae separated by the anterior lobe of the metasternal plate, which is 
never linear. between the coxae. 2 =~ 22242 3254 -- 6 see ec bese eee 5 


1 Odontaeus is asynonym. See Introduction and also discussion under Bradycinetulus. 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 97 


Bp liivtralwith: 5, discal striagou..5) 21 s2steleen se eeck ue leeeecsule Eucanthus 
avira with discal siriaglas uel. vio pie euscels Juss deen ok Bolbelasmus 

5. Base of elytia margined, metasternal plate pyriform in outline, pronotum 
without postapical carina, first elytral stria interrupted by scutellum____ 6 

Base of elytra not margined, metasternal plate rhomboid in outline, pronotum 
with postapical carina, first two elytral striae interrupted by scutellum. 
Bolborhombus, new genus 

6. Apex of tibia of middle and hind legs deeply emarginate on outer side, the 
angle adjacent to spurs appearing almost as a fixed spur; without prosternal 
pe seth She tee eto) es tet ert eaaathes Bradycinetulus 
Apex of tibia of middle and hind legs obliquely truncate; prosternal spine 
behind anterior coxae transverse, doubly pointed, and remote from acutely 
aneied. intercoxal. piece... 522.2522. Bolbocerastes, new genus 


Genus Bolbelasmus Boucomont, 1911 


Bolbelasmus Boucomont, Ann. Soc. Ent. France, vol. 79 (1910), p. 835, 1911. 
Kolbeus Boucomont, Ann. Soc. Ent. France, vol. 79 (1910), p. 335, 1911.-—Chapin, 
Proc. Biol. Soc. Washington, vol. 59, p. 79, 1946. 


Chapin, 1946, placed Bradycinetus hornii Rivers and B. minor 
Linell in the genus Kolbeus, basing his action on their close relation- 
ship to Kolbeus arcuatus (Bates). Examination of specimens of these 
species in comparison with Bolbelasmus gallicus (Mulsant) and B. 
unicorne (Shrank), and in the light of Boucomont’s key to the genera, 
places B. minor Linell and B. hornii Rivers in Bolbelasmus rather 
than in Kolbeus. 

The genus Kolbeus was erected without a genotype by Boucomont, 
1911, for the species Bolboceras arcuatus Bates and B. coreanus Kolbe. 
Lucas, 1920 (Catalogus . . . Coleopterorum, p. 356), selected 
B. coreanus Kolbe, a species found in Korea and China, as the 
genotype. The species B. arcuatus Bates almost exactly duplicates 
Bolbelasmus minor (Linell) except for the slightly narrower scutellum 
and lack of a marginal line over the middle half of the base of the 
pronotum. Since a direct comparison of specimens shows such minor 
differences, I transfer arcuatus Bates to the genus Bolbelasmus. 
Whether the genus Kolbeus should be synonymized with Bolbelasmus 
will depend upon an examination of its genotype, a species not avail- 
able for study at present. Further basis for placing arcuatus in Bol- 
belasmus is found in similarity of the aedeagi (see fig. 14). 

Genotype.—Scarabaeus gallicus Mulsant, Histoire naturelle des 
Coléoptéres de France, Lamellicornes, p. 350, 1842. (Present desig- 
nation. ) 

Key to the species of Bolbelasmus 
1. Base of pronotum not margined over middle half; length 7 to 9 mm. (Cali- 


Ronny WVEGsico) st f- Saeer ote ro Bseue sh. folie arcuatus (Bates) 
Base of pronotum completely margined____________.._..-------------- 2 


98 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


2. Elytral striae coarsely, deeply punctate asin Hucanthus; genae obtusely rounded; 
surface of head and clypeus similarly densely, coarsely punctate in male; 
frontal carina weakly tri-tuberculate in female; length 7 to 10 mm. (Texas). 

minor (Linell) 

Elytral striae finely to moderately punctate; genae angulate externally; 
surface of clypeus roughly, densely, coarsely punctate, remaining surface of 
head much smoother and more finely punctate in the male; frontal carina 
not trinodose in female; length 14 mm. (California)____-___ hornii (Rivers) 


Bolbelasmus arcuatus (Bates), new combination 


Ficure 14, | 


Bolboceras arcuatus Bates, Biologia Centrali-Americana, Coleoptera, vol. 2, pt. 2, 
Bott Sei 

Kolbeus arcuatus Boucomont, Ann. Soc. Ent. France, vol. 79 (1910), p. 336, 1911.— 
Chapin, Proc. Biol. Soc. Washington, vol. 59, p. 79, 1946. 


Bolbelasmus arcuatus (Bates) is a small, dark red-brown, shining 
species, 7 to 9 mm. in length, readily recognized through the lack of 
basal margins on both pronotum and elytra. The labrum is widely, 
shallowly emarginate in front; the clypeus is almost evenly arcuate 
from side to side; the clypeal suture is distinct; the anterior angles of 
the pronotum are sharply right angled; the scutellum is elongate; 
the strial punctures are coarse, in the outside row at the apex very coarse 
and unusually deep, practically perforating the elytra; the elytra 
have a row of widely spaced, long, bristlelike hairs near the margin; 
the male has a single median conical cephalic horn and a low, wide, 
median thoracic prominence and anterior shallow depression on each 
side; the female has a wide, more-or-less tri-tuberculate carina on the 
head and a simple, fine, sharp, transverse carina on the pronotum. 

Type.—Type probably among the Biologia specimens in the British 
Museum (Natural History). 

Type locality —Not designated among the localities originally listed, 
“Mexico, Cordova, Toxpan, Playa Vicente (Sallé), Colima City 
(Hége) ; Nicaragua, Chontales (Belt, Janson).”’ 

Specimens examined.—19. 

Distribution —Costa Rica: Tilarén (Guanacaste), San Pedro de 
Montes de Oca, San José, and *Dominical. Guatemala. México: 
Colima, El Sabino, Compostela, Mazatlén, Monterey, *Monclova, 
and *Guadalajara. United States: a single specimen labeled ‘Los 
Angeles, California, H. Klages Coll.,”’ in the Carnegie Museum, 
Pittsburgh, Pa. 


Ficure 14.—Ventral and lateral views of the aedeagi of Bradycinetulus and closely related 
species: a-c, Species of Bolborhombus: a, parvulus, new species; b, angulus (Robinson); 
c, schaeffert (Boucomont). d-g, Species of Bolbocerastes: d, peninsularis (Schaeffer); ¢, 
regalis, new species;| f, serratus (LeConte); g, imperialis, new species. h-j, Species of 
Bradycinetulus: h, ferrugineus (Palisot de Beauvois); 1, rex, new species; j, fossatus 
(Haldeman). &k-n, Species of Bolbelasmus: k, gallicus (Mulsant); 1, arcuatus (Bates); 
m, hornit (Rivers); 2, minor (Linell). 


100 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Bolbelasmus minor (Linell), new combination 


Ficure 14, n 


Bradycinetus minor Linell, Proc. U. 8. Nat. Mus., vol. 18, p. 723, 1895. 
Bolboceras minor Schaeffer, Trans. Amer. Ent. Soc., vol. 32, p. 253, 1906. 
Kolbeus minor Chapin, Proc. Biol. Soc. Washington, vol. 59, p. 79, 1946. 

Bolbelasmus minor (Linell) almost exactly duplicates B. arcuatus 
(Bates) except that it averages slightly larger, the punctures are 
coarser throughout, and the pronotum is completely and distinctly 
margined basally. It is a small, dark red-brown, shining species, 
7 to 10 mm. in length; the labrum is widely, shallowly emarginate in 
front; the clypeus is almost evenly arcuate from side to side; the 
clypeal suture is distinct; the anterior angles of the pronotum are 
sharply right angled; the scutellum is elongate; the strial punctures 
are very coarse, those at the apex of the elytra being tubular and 
so deep as to nearly perforate the elytra; near the margin of the elytra 
is a row of widely spaced, long, bristlelike hairs; the male has the 
head and clypeus similarly densely punctate, the head bears a median, 
conical horn, and the pronotum has a low, wide, median prominence 
with anterior, shallow depressions each side; the female has a wide, 
transverse, tri-tuberculate carina on the head and a simple, fine, 
transverse carina on the pronotum. 

Type.—Holotype female, USNM 562. 

Type locality —San Diego, Tex. 

Specimens examined.—21. 

Distribution.—Texas: Hidalgo County, Weslaco, Brownsville, Kings- 
ville, and San Diego. 

Season.—May 16 to June 16. 


Bolbelasmus hornii (Rivers), new combination 


Ficure 14, m 


Bradycinetus hornii Rivers, Bull. California Acad. Sci., vol. 2, p. 61, 1886. 
Bolboceras hornit Schaeffer, Trans. Amer. Ent. Soc., vol. 32, p. 253, 1906. 
Bradycinetulus hornii Cockerell, Ent. News, vol. 17, p. 242, 1906. 

Kolbeus hornii Chapin, Proc. Biol. Soc. Washington, vol. 59, p. 79, 1946. 


Bolbelasmus hornii (Rivers) is quite similar to B. minor (Linell) but 
is considerably larger, lighter in color, and has much finer strial 
punctures. The scutellum is not so elongate, being nearly an equi- 
lateral triangle with arcuate sides. Other characters given in the 
key should readily identify this species. 

Type.—Location of type unknown to me. (According to Dr. 
VanDyke the entire Rivers Collection was purchased by Dr. Walther 
Horn and it is presumably in the Berlin-Dahlem collection.) 

Type locality—‘Sonora, Tuolumne Co., Cal. Found also in 
Sacramento Co.” 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 10] 


Specimens examined.—349. 

Distribution.—California: Sequoia National Park, Napa, Kaweah, 
Camp Greely (Tulare County), Mendocino, Mokelumne Hill, Camp 
Greely (Fresno County), Three Rivers, El Dorado County, Coulter- 
ville, Sonoma County, Potwisha, San Joaquin Experiment Station 
(Madera County), Mount Diablo, Sacramento, Mineralking, La Jon 
Pass, Rattlesnake (Placer County), *Calaveras County, and *O’Neals 
(Madera County). Baja California: Hamilton Ranch. 

Season.—January 19 to May 8, with greatest number in April. 


Genus Bradycinetulus Cockerell, 1906 


Bolboceras Kirby (in part), Trans, Linn. Soc. London, vol. 12, pt. 2, p. 459, 1818. 
(Type not designated.) 

Amechanus Horn (not Thomson, 1864), Trans. Amer. Ent. Soc., vol. 3, p. 48, 
1870. 

Bradycinetus Horn (not Sars, 1865), Trans. Amer. Ent. Soc., vol. 3, p. 334, 1871. 

Bradycellus Schaeffer (not Erichson, 1837), Trans. Amer. Ent. Soc., vol. 32, p. 

249, 1906. 
Bradycinetulus Cockerell, Ent. News, vol. 17, p. 242, 1906. (Type not designated.) 


The genus Bolboceras was erected by Kirby in 1818 for the eight 
species Scarabaeus mobilicornis Linnaeus, S. mobilicornis var. tes- 
taceus Fabricius, S. qguadridens Fabricius, S. farctus Fabricius, S. 
lazarus Fabricius, S. cyclops Olivier, S. cephus Fabricius, and Bol- 
boceras australasiae Kirby. Technically no genotype was designated 
although he wrote, ‘“My details of Bolboceras were taken from B. 
quadridens.”” Nearly all of the species mentioned have since been 
moved to other genera. Curtis, 1829 (British Ent., vol. 1, pt. 1, p. 
74), selected the species Scarabaeus mobilicornis Fabricius as the type 
of Bolboceras. Therefore, since our species of Odontaeus are con- 
generic with mobilicornis, they now take the generic name Bolboceras 
and it becomes necessary to find an available name for those species 
we have formerly placed in Bolboceras. A search of the literature 
indicates that name must be Bradycinetulus, as shown in the 
synonymy. 

Characters for Bradycinetulus, in addition to those listed in the key 
to genera, include the following: The mandibles are evenly arcuate ex- 
ternally; the median prominence of the pronotum is developed into a 
pair of horns in the male; the scutellum is wider than long and dis- 
tinctly punctate with fine to moderate punctures; and the posterior 
vertical face of the prosternal intercoxal piece is wide and flat, with 
the ventral edge evenly arcuate or slightly angulate at middle. The 
three species placed here are all large, 17 to 21 mm. in length by 10 
to 12mm.in width. They are sufficiently differentiated in the follow- 
ing key. 

Genotype —Scarabaeus ferrugineus Palisot de Beauvois, 1809. (Pres- 
ent designation.) 


102 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Key to the species of Bradycinetulus 


1. Eye canthus nearly straight edged laterally, wide clypeal horn of male bisinuate 
apically, the external angles projecting forward, acute, and scarcely thicker 
than the adjacent edges, which converge posteriorly; median thoracic horns 
heavy, directed upward, their sharply rounded tips recurved posteriorly 
(North Carolina to Florida and Mississippi). 

ferrugineus (Palisot de Beauvois) 
Kye canthus notched or emarginate, not straight edged laterally___________ 2 

2. Wide clypeal horn of male distinctly trinodose apically, the external angles 
not projecting beyond median point, lateral edges parallel; median thoracic 
horns heavy, projecting forward and outward, sharply rounded and very 
little recurved (Nebraska to Texas and Arkansas)___-_fossatus (Haldeman) 

Apex of clypeal horn of male widely truncate, straight across, the external 
angles slightly nodose, lateral edges parallel over apical two-fifths, then 
diverging to base; median thoracic horns slender, directed forward and up- 
ward, basally flattened in front and behind (Texas)____rex, new species. 


Bradycinetulus ferrugineus (Palisot de Beauvois), 1809 
Ficure 14, h: Puats 3 


Scarabaeus ferrugineus Palisot de Beauvois, Insectes recueillis en Afrique et en 
Amérique, livr. 6, p. 90, 1809. 

Bolboceras lecontei Dejean, Catalogue des Coléoptéres . . ., ed. 3, vol. 3, p. 149, 
1833. 

Athyreus ferrugineus Klug, Abh. Berlin Acad., 1843, p. 22. 

Bolboceras ferrugineus Lacordaire, Histoire naturelle des insectes, vol. 3, p. 148, 
1856. 

Amechanus ferrugineus Horn, Trans. Amer. Ent. Soc., vol. 3, p. 48, 1870. 

Bradycinetus ferrugineus Horn, Ent. Amer., vol. 1, p. 89, 1885. 

Athyreus (Bradicinetus) ferrugineus Boucomont, in Wytsmann, Genera insectorum, 
fase. 7, p. 8, 1902. 

Bradycinetus ferrugineus Schaeffer, Trans. Amer. Ent. Soc., vol. 32, p. 250, 1906. 

Bradycinetulus ferrugineus Cockerell, Ent. News, vol. 17, p. 242, 1906. 

Bolboceras (Amechamus) ferrugineus Boucomont, Ann. Soc. Ent. France, vol. 79, 
p. 341, 1910. 


Type.—Location of type unknown to me. 

Type locality —South Carolina. 

Specimens exramined.—107. 

Distribution.—North Carolina: West End, Southern Pines, **Beau- 
fort, *Hamlet, *Carolina Beach, and **Tarboro. South Carolina: 
Meredith, Florence, Windsor, Johns Island, and Bulls Island. Geor- 
gia: Bainbridge, Spring Creek (Decatur County), and Augusta. Flor- 
ida: Port Saint Joe, Kissimmi, Orlando, Sanford, Enterprise, Miami, 
Crescent City, Indian River, Cedar Keys, *Lutz, *Wacissa, *Gaines- 
ville, **Saint Augustine, **LaGrange, and **Dunedin. Alabama: 
**Lee County, and **Mobile County. Mississippi: Ocean Springs, 
Biloxi, and Leakesville. 

Season.—March 11 to September 23, with the largest number of 
records in June and July. 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 103 


Bradycinetulus fossatus (Haldeman), 1853 
Ficure 14, j; PLats 3 


Bolboceras fossatus Haldeman, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 362, 
1853. 

Amechanus fossatus Horn, Trans. Amer. Ent. Soc., vol. 3, p. 48, 1870. 

Bradycinetus fossatus Schaeffer, Trans. Amer. Ent. Soc., vol. 32, p. 250, 1906. 

Bradycinetulus fossator Cockerell, Ent. News, vol. 17, p. 242, 1906. 

Bolboceras (Amechamus) fossatus Boucomont, Ann. Soc. Ent. France, vol. 79, 
p. 341, 1910. 

In a few male specimens of Bradycinetulus fossatus (Haldeman) the 
outer cariniform edges bordering the thoracic fossae show a slight 
tendency to break near the upper end to form a second inner carina, a 
step toward the condition found in Bolbocerastes. 

Type.—In LeConte collection, MCZ. 

Type locality —Texas. 

Specimens examined.—102. 

Distribution.—Nebraska: Imperial. Kansas: Reno County, Me- 
dora, and Sylvia. Oklahoma: Noble County, Cleveland County, 
Payne County, Thomas, Cleo Springs, Ferris, and *Alva. Arkansas: 
Pine Bluff. Texas: Fedor, Henrietta, Tyler, Jacksonville, Gold- 
thwaite, Robinson, Lexington, Jefferson, College Station, Lindale, 
Bexar County, Dallas, Victoria, Columbus, Paris, Austin, San Antonio, 
and Morris County. 

Season.—May 16 to August, with most records in June. 


Bradycinetulus rex, new species 


Figure 14, 7; PLats 3 


Holotype male, length 21 mm.,width 13mm. In general appearance 
very similar to Bradycinetulus ferrugineus (Palisot de Beauvois) and 
B. fossatus (Haldeman), rufo-testaceous, moderately shining, densely 
hairy beneath. Clypeus perpendicular in front, the nearly flat, verti- 
cal face extending upward to form a broad truncate horn about one- 
fourth wider than high. The face and sides of the horn reticulate, 
the enclosed spaces concave, shallow, and coarse; the truncate apex 
scarcely thickened, with the external angles very slightly nodose; 
the sides parallel over apical two-fifths, diverging posteriorly ; margin 
thin, irregularly serrated, elevated just before the eye canthus to a 
very moderate right-angled prominence, the posterior edge of which 
continues posteriorly as a low, sharp carina separating the eye canthus 
from the flattened discal surface of the head. Canthus sloping away 
from the discal level of head; anterior and lateral margins sinuate to 


232890—53——2 


104 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


the rounded, elevated, anterior angle, surface ridged inward from the 
angle, depressed along anterior edge and in posterior angle. Margin 
of head along inner edge of the eye with a reflexed carina, which grad- 
ually curves away from the eye, ‘increases to a maximum height, then 
angles sharply inward and backward and sharply descends to general 
level of head ; head surface, including eye canthi, reticulate-tuberculate- 
punctate, very rough anteriorly on the concave posterior of the clypeal 
horn, gradually smoother over discal area, which is bounded poste- 
riorly by a low, rounded ridge between the eyes; the reticulation and 
tubercles of the discal area flatten until the outlines of the coarse, 
shallow punctures are barely traceable; head behind the ridge smooth 
at middle but gradually roughened near the eyes. 

Pronotum similar to that of B. ferrugineus (Palisot de Beauvois) 
but with the horns of the median prominence and the high lateral 
edges of the depressions much farther forward. ‘The anterior edge 
of the pronotum very finely margined, the middle third arching slightly 
over the base of the head, each outer third without a curve, straight, 
diverging anteriorly to the right-angled anterior angles, slightly 
concave close along these straight edges, then very obtusely convex 
and rounding into the deep lateral depressions, the middle third with 
a smaller, closer, parallel convexity and a moderate concavity at each 
end; pronotum laterally with two very large deep fossae on each side, 
the lower wider and less deep, widely sinuating the lateral margin 
from middle to near the anterior angles, the upper very deep, the 
outer edge thrown up in a high, thin, curving edge which descends 
to the posterior boundary of the lower cavity; posterior angles distinct 
but obtusely rounded; sides and base margined, the edge rather finely 
serrate laterally; pronotum convex basally; disc forming a broad 
median prominence with a sharp horn pointing upward on each side, 
the horns being rather sharply undercut below and basally flattened 
in front and behind; a deep median line from base forward over the 
median prominence and down between the horns. Surface of the 
pronotum alutaceous and finely scabrous back of the anterior margin, 
especially at the sides, then shallowly, coarsely punctate over the 
inner lower surface of the lateral fossae; posterior angles similarly 
finely alutaceous and closely, coarsely, shallowly punctate; elsewhere 
finely to minutely, sparsely punctate throughout, except for the very 
smooth surface of the deeper fossae; and with a large group of close 
very coarse punctures on each side back of the horn and above the 
deeper fossa, another group following the median line and spreading 
anteriorly to bases of the horns. 

scutellum finely punctate. Elytra not strongly shining; minutely 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 105 


alutaceous under high magnification; margined at base; striae mod- 
erately impressed, the strial punctures fine, moderately deep; first 
stria interrupted at scutellum; the second indistinct near scutellum. 

Metasternal lobe separating the middle coxae half as wide as the 
plate behind the coxae; lateral edges sharply carinate. 

Female unknown. 

Type.—Holotype male, USNM 61455. 

Type locality—Sarita, Tex. Holotype collected on bare sand, 
Novy. 30, 1911. 

Paratypes——One male, CAS, Corpus Christi, Tex., June 28, 1942, 
E. 5S. Ross; one male, in Mark Robinson collection, Kingsville, Tex., 
C. T. Reed. 

Remarks.—Bradycinetulus rex may be readily separated from the 
two closely allied species B. ferrugineus (Palisot de Beauvois) and B. 
fossatus (Haldeman) by the obvious differences in the clypeal and 
thoracic horns. It may be noted also that the last two species have 
only a single deep fossa on each side of the pronotum, while in B. rez 
this is doubled, the lower cavity sinuating the lateral margin. 


Bolbocerastes, new genus 


This genus is closely allied to Bradycinetulus but differs in that the 
apex of the tibia of the midleg and hind leg is obliquely truncate, the 
scutellum is smooth or minutely punctate, the mandibles are parallel, 
nearly straight sided and bent sharply inward anteriorly, the sides and 
anterior edges forming a rectangle, the median prominence of the 
pronotum is without horns, two elevated carinae are present on each 
side of the pronotum, the prosternal spine behind the anterior coxae 
is transverse, doubly pointed and remote from the acutely angled 
intercoxal piece, and the aedeagus of the male is of a characteristic form 
(see fig. 14). 

Genotype.—Bolbocerastes regalis, new species. 


Key to the species of Bolbocerastes 


1. Anterior clypeal horn of male widely truncate; anterior and median horns 
of female usually equal in width, smooth back of median horn (Arizona, 
NevadariOalifemiia) 413. seal bea tety sd beaut Avie: regalis, new species 

Anterior clypeal horn of male narrowly truncate or rounded; anterior horn 
narrower than median horn in female, punctate back of median horn. 2 

2. Dull, dark reddish brown; elytral striae extremely fine, not impressed, the 
intervals completely flat even in reflected light, strial punctures very fine; 
fully developed male with the anterior clypeal horn narrowly rounded and 
very little advanced, much closer and more nearly in line with the strongly 
developed lateral horns than in serratus and imperialis (California, Baja 
CST ONINIS 2 ea ee Pe ne eae em OPE eo peninsularis (Schaeffer) 

Elytral striae fine to moderate, usually at least slightly impressed, the intervals 
weakly convex in reflected light; fully developed males not as above__. 3 


106 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


3. Lateral serrations of pronotum arising between the two edges of a doubly 
carinate margin; elytral striae moderately impressed, strial punctures dis- 
tinct; color reddish brown, usually quite strongly shining; tip of clypeal 
horn rounded in male, inner thoracic carina without secondary angle at 
upper end; female cephalic horn rarely higher than clypeal horn, carina 
extending back from clypeal horn on each side sharp, strong, usually dark 
colored, lateral secondary horns nearer midway between clypeal and cephalic 
horns than to the cephalic horn, discal area of head distinctly punctate 
throughout, fine punctures of median thoracic prominence distinct and 
noticeable (Kansas, Oklahoma, Texas, New Mexico)-serratus (LeConte) 

Pronotum without or with only traces of a doubly carinate margin laterally, 
the serrations appearing as thickenings of a single sharp edge; well developed 
males with a secondary angle at upper end of inner thoracic carina_.__ 4 

4. Elytral striae much less distinct, the elytra smoother in appearance, nearly 
as in peninsularis, color lighter, more yellow-brown, less strongly shining; 
fully developed male with tip of clypeal horn narrowly truncate and well 
forward from the lateral horns, upper lateral thoracic carina of the male 
frequently with a noticeable secondary angle at upper end; female median 
cephalic horn often very high, usually higher than clypeal horn, carina 
extending back to lateral secondary horn weak, sometimes obliterated, not 
darker in color, the lateral secondary horns weak, usually much farther 
back, more nearly in line with the cephalic horn, discal area of head usually 
with only scattered punctures behind median horn (Baja California, Cali- 


fornia, Arizona, western Texas). —-_-_-__-.-----= imperialis, new species 
Elytral striae deeper, color red-brown, more strongly shining as in serratus, 
otherwise as above (Kansas) - ------ imperialis kansanus, new subspecies 


Bolbocerastes regalis, new species 


Figure 14, e; Puats 4 


Holotype male, length 18 mm., width 11.5 mm. In general appear- 
ance closely similar to Bolbocerastes serratus (LeConte), ferrugineus, 
shining, densely hairy beneath. Clypeus perpendicular in front, the 
flat vertical face extending upward to form a broad truncate horn, 
the vertical height subequal to base, the lateral edges arcuate from 
base to upper edge, narrowing at basal third to half the basal width 
then expanding to three-fourths, the upper edge weakly emarginate 
and subparallel with base, shining black in color, rounded and knoblike 
at each end; the sharply defined lateral triangular faces of the horn 
concave and posteriorly elevated from base into secondary horns half 
as high; posteriorly the broad curving slope of the clypeal horn merges 
into the flattened upper surface of the head, this quadrangular surface 
with a secondary horn at each anterior angle and with sharply defined 
carinalike subparallel lateral margins from which the prominent, 
widely explanate eye canthus slopes downward and outward. Occip- 
ital area smooth; gradually, finely, closely, shallowly punctate 
anteriorly; anterior and lateral faces of clypeal horn, labrum, eye 
canthi, and upper surface of mandibles finely reticulate-scabrous. 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 107 


Broadly rounded labrum twice as wide as long. Upper surface of 
mandibles concave, sides straight, bent at right angles in front. 

Pronotum retuse in front; the deeper lateral impressions extending 
upward on each side of the median prominence and bounded laterally 
by two carinae on each side; the lower external carina starting near 
the serrate lateral thoracic margin and extending upward parallel 
with the basal thoracic margin for a distance equal to half the length 
of the lateral margin; the upper carina starting inside the upper end 
of the lower and extending upward for a slightly shorter distance; 
viewed from the side the lower carina is nearly straight, the upper 
end with a gentle downward curve, the lower end with a noticeable 
but low toothlike elevation; in outline the upper carina is half as 
high as long, sloping upward from a sharp basal curve to a rounded 
right-angled peak, then down on a concave curve to a noticeable sec- 
ondary angle. The median pronotal prominence sharply declivous in 
front, upper surface divided by an anteriorly increasingly depressed 
median line, the parallel sides with sharply rounded edges anteriorly; 
retuse front impunctate except for a subapical line of close, weak-to- 
obsolete punctures; median prominence with fine, close punctures 
shading to minute punctation across base; a few obsolete moderate 
punctures in the shallow lateral depressions outside the carinae; basal 
bead distinct. Lateral edges strongly serrate, as in Bolbocerastes 
serratus (LeConte). 

Elytra not strongly shining; striae scarcely impressed; strial 
punctures fine, moderately close, intervals practically flat, very 
minutely punctulate. 

Metasternal lobe separating middle coxae half as wide as plate 
behind the coxae. 

Female allotype, length 21 mm., width 10 mm. Similar to the male 
except that the thoracic prominences are much less developed and 
the head is different. The vertical face of the clypeus is only half as 
high as the basal length, the upper edge less than half the basal length, 
and from each end a curving carina extends outward to top of the 
secondary horn or tubercle; from the flattened area back of and about 
twice as high as the secondary tubercles arises a widely truncate 
median horn; anterior to this median horn the surface is reticulate- 
scabrous, posteriorly it is smooth. 

Type.—Holotype male, USNM 61076. 

Type locality —The holotype bears the following label data: ‘3206. 
Colo. R. bottom. Monument 204. Mex. Bd. line Mar. 20-31, ’94. 
USNM Ace. No. 28133 Dr. E. A. Mearns.”” Monument 204 is approx- 
imately 20 miles south of Yuma, Ariz., near San Luis, México, its 
exact location being lat. 32° 29’ 03.59’’, long. 114° 46’ 44.797’. 


108 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Allotype and paratypes——Allotype and 97 paratypes bear data as 
follows: Nevada: 1, Hawthorn, June 8, 1943, Wallace; 1, Mount 
Charleston, 8,000—10,000 feet, Aug. 6, 1944, Zinn. | 

California: 2, Baker, Mar. 27, 1935, and April 1935, Saylor; 1, 
California (State label only); 3, Coachella, May 12, 13, 17, 1927, Van 
Duzee; 3, Coachella, Riverside County, Apr. 28, 1928, May 9, 1927, 
July 15, 1927, Wymore; 2, Coachella Valley, May 9, 1927, Van Dyke, 
Aug. 20, 1949, v. d. Bosch; 1, Coyote Wells, Imperial County, June 5, 
1949, Hurd; 1, Essex, Apr. 29, 1937; 1, Holtville, June 28, 1936, 
Cazier; 14, Imperial County, Carrizo Light, Mar. 14, 17, 18, 22, 1928, 
Searl; 1, Indian Wells, Riverside County, Apr. 25, 1939, Martin; 
8, junction of Highways 78 and 99, Imperial County, Apr. 15, 1944, 
Tieman; 4, Mecca, Mar. 25, 1922, Benedict; 3, Murrays Dam, San 
Diego County, Cottle; 1, Needles, Apr. 1, 1941, Linsley; 2, Palm 
Springs, May 20, 1905, May 9, 1932; 2, San Clemente Island, Apr. 2, 3, 
1939, Channel Islands Biological Survey, Von Bloeker; 1, San Diego 
County, Mar. 10, 1928, light, Well; 7, Yermo, San Bernardino County, 
Apr. 11, 1936, Comstock, Sept. 6-30, 1939, Pierce. 

Arizona: 7, Arizona (State label only); 4, Ehrenberg, Oct. 18, 1938, 
Mar. 21, 23, 30, 1939, Parker; 6 (includes allotype), Gila Bend, Mar. 1, 
1930, Lebart; 1, Holbrook, May 5, 1941, Dukes; 1, Navaho County, 
Duncan; 1, Phoenix; 1, Red Rock, Spring 1930, Burdett; 1, Tucson, 
April 192—, Duncan; 11, Yuma, February 1910, Carlson, Mar. 12, 18, 
1912, Slevin, May 15, 1939, Aitken. 

México, Sonora: 1, Choya Bay, Mar. 27, 1949, Bradt; 1, 30 miles 
southwest of Sonoyta, 500 feet, Mar. 31, 1949, Bradt; 4, Puerto 
Libertad, Mar. 15, 1939, Parker. 

Paratypes in AMNH, CAS, LAC, KAN, MCZ, CAL, CU, OSd, 
MICH, INHS, USNM, and the private collections of Mark Robinson, 
_Henry Howden, C. A. Frost, A. T. McClay, F. H. Parker, D. K. 
Duncan, and R. H. Arnett. 

Remarks.—Bolbocerastes regalis varies from 16 to 20 mm. in length 
and from 10 to 12.5 mm. in width. It is closely similar to B. serratus 
(LeConte) but the males differ noticeably in having a higher, wide, 
truncate clypeal horn, smoother, less punctate pronotum, and on each 
side a doubly angulate inner pronotal carina which is always higher 
in the middle. In B. serratus (LeConte) the anterior clypeal horn is 
bluntly rounded, the pronotal punctures are more evident throughout, 
and the inner lateral pronotal carina is not doubly angulate or higher 
in the middle. The truncated tops of the anterior and median horns 
are about equal in width in females of B. regalis and the whole anterior 
part of the head, including clypeal and cephalic horns, appears higher 
than the posterior part, like an elevated snout. The lateral horns are 
most prominent in this species. 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 109 


Bolbocerastes imperialis, new species 


Figure 14, g; PLate 4 


Holotype male, length 19 mm., width 11 mm. Very similar to 
Bolbocerastes serratus (LeConte); ferrugineus; moderately shining; 
hairy beneath. Vertical face of clypeal horn inclined forward about 
30 degrees from perpendicular, slightly convex when viewed from side, 
height seven-tenths width at base, width of the truncate apex slightly 
less than half the basal width, the apex arcuate posteriorly at 
each side; the posterior face of the horn concave from side to side 
between the weakly carinate edges, which converge very slightly from 
apex to base. General level of the head slightly more than half the 
height of the clypeal horn above the wide, truncate labrum; this flat- 
tened area sloping upward posteriorly and bounded by a ridge or 
carina deeply arcuate posteriorly from the anterior edge of one eye to 
the other and continuing forward on each side to the tips of the weakly 
elevated, right-angled, secondary horns, then converging to the an- 
terior edge of the clypeal horn and forming the upper edge of the tri- 
angular lateral area at base of the horn; flattened area of head slightly 
depressed laterally above the prominent quadrangular eye canthus, 
which slopes arcuately downward, outward, and finally upward at the 
outer angles; entire surface in front of the curving posterior ridge 
densely, roughly punctate, smooth back of the ridge. Mandibles 
concave dorsally; strongly angled front and back. 

Pronotum retuse in front and with serrate lateral margins as in B. 
serratus (LeConte) and B. regalis; the median prominence and lateral 
carinae nearly as in B. regalis but with the outer carina with the upper 
end curving slightly away and not quite parallel with the basal mar- 
gin when viewed from the side; the inner lateral carina, viewed from 
the side, with the lower end rising abruptly to a right angle, then 
sloping evenly upward to a right-angled peak at middle, then down 
in a strongly sinuate in-and-out curve; both carinae, viewed from 
directly in front, evenly arcuate, with the peak angle of the inner carina 
strongly bent inward; the median prominence slightly constricted 
toward the base, with the sides weakly angulate in front of the con- 
striction and the surface more broadly and less deeply concave from 
side to side than in B. regalis and evenly arcuate from front to back; a 
distinct median line evident basally; the apical and deeper lateral 
pronotal depressions extending upward between the lateral carinae; 
the median prominence impunctate except for half a dozen or so close, 
moderate punctures on each side at base of the vertical sinus; above 
these about twenty or so scattered fine punctures on the inner slope 
of the vertical sinus, and a very few fine punctures indicating a sub- 
apical line; median prominence and lateral basal area back of the 


110 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


carinae closely, finely punctate; smoother and more finely punctate 
across base at middle; groups of moderate punctures in the depressions 
back of the carinae and an irregular row of close moderate punctures 
just inside the basal bead. 

Scutellum closely, minutely punctate. Elytra one-sixth wider than 
long; extremely, finely alutaceous under moderate magnification; 
striae scarcely impressed; strial punctures fine, distant about four 
diameters; intervals with very minute scattered punctures. 

Metasternal lobe separating middle coxae one-third as wide as the 
plate behind the coxae. 

Allotype female, length 14 mm., width 9mm. Similar to the male 
except that the pronotal prominences are less strongly developed and 
the head is different. The vertical face of the clypeus is one-third 
as high as basal length; its upper truncate edge also one-third the 
length of the base; the concave triangular sides of the clypeal horn 
with the upper edge about twice as long as the anterior edge; the 
wide truncate frontal horn is about three-fourths as high as wide and 
about four times as high as the low secondary lateral horns, or tubercles. 

Type.—Holotype male, USNM 61077. 

Type locality.—Imperial County, Calif., “‘on the Experiment Farm,” 
June 1912, J. C. Bridwell. 

Allotype and paratypes.—Allotype female and 210 paratypes bear 
the following data: Texas: 1, Castolon, June 1, 1928, Bibby; 1, 
Tornilla Flat, Big Bend National Park, July 12, 1948, desert, 3,650 
feet, at light, Werner and Nutting; 2, Boquillas, Brewster County, 
July 7, 1948, C. and P. Vaurie; 1, Marathon, June 14, 1948, D. Rocke- 
feller Expedition, Cazier; 1, Reeves County, June 10, 1939, Robinson. 

New Mexico: 1, Artesia, July 29, 1937, D. J. and J. N. Knull; 1, 
White Sands, July 23, 1933, Benedict. 

Arizona: 4, Agua Fria, Aug. 6, 1917, Bequaert, June 26, 1938, 
Duncan; 2, Aquila, Maricopa County, Aug. 21, 22, 1927, CU lot 542, 
sub 330; 2, Arlington, July 25, 1948, Anderson; 15, Arizona (State 
label only); 5, Avondale Ranch, Agua Fria River, Aug. 7, 1917, 
Wheeler leg.; 4, Baboquivari Mountains, July 24, 1941, Beamer, 
July 24, 1941, Todd; 2, Buckeye, July 30, 31, 1935, at light, Christen- 
son; 1, Cashion, Maricopa County, Aug. 25, 1949, at light, Werner 
and Nutting; 5, Cibola, August 1911; 1, Continental, Pima County, 
July 29, 1948, Ball; 1, Coyote Mountains, Aug. 4, 7, 1916; 1, Dome, 
July 21, 1924, Van Duzee; 4, Ehrenberg, July 20, 1946, June 12, 
1943, Grassman, July 18, 1939, Stager, June 21, 1938, Parker; 10, 
Globe, Duncan, Sept. 8, 1930, July 11, 1931, July, August, Duncan 
and Parker; 7, Hope, Yuma County, Aug. 12, 1948, greasewood desert, 
1,400 feet, at light, Werner and Nutting; 1, Hot Springs, July 16 
(year not given), at light, Morrill; 2, Huachuca Mountains; 2, Kits 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 111 


Peak, Rincon, Baboquivari Mountains, Aug. 1, 4, 1916, about 1,050 
feet; 1, Littlefield; 1, Marinette, Aug. 4, 1918, Schiffel; 1, Olberg, 
Sept. 13, 1936, Crandall; 1, Palmerlee; 8, Phinax (sic), September 
and November 1921; 17, Phoenix, Sept. 12, 1935, Crandell, May 12, 
1934, Parker, May 29, 1922, Odell, Aug. 2, 1917, CU Biol. Expedition, 
May 17, 1944, Parker, Apr. 29, 1941, Parker, May 7, 1918, Bradley, 
Sept. 12, 1926; 1, Pima County, Sept. 1, 1925, Marsh; 1, Prescott; 2, 
Sacoton, Gilman; 1, Safford, September (no date given), Duncan; 
2, San Bernardino Ranch, Cochise County, 3,600 feet, sycamore- 
willow in valley at light, July 25, 1949, Werner and Nutting, July 16, 
1949, light, Parker; 1, San Luis, Yuma County, Aug. 11, 1940; 1, 
Talklai, Pollock; 20, Tucson, July 21 (no year given) Hubbard and 
Schwarz, July 30, 1937, Parker, August 1912, Breitenbecker, Aug. 20, 
1932, Flock, August 1935, Wharton, October 1927, Hamilton (2,500 
feet), July 10, 1950, Aug. 8, 1950, Bradt, Oct. 14, 1919; 3, Wellton, 
June 13, 1939, at light, Stitt; 1, Welton, Aug. 9, 1917, Wheeler leg.; 
1, Yuma. 

California: 7, Blythe, May 19, 1942, Allen, Aug. 26, Hurd, April 
16, 1945, Constantine, May 19, 1943, Vargas, July 22, 1947, July 8, 
1947; 4, Blythe, Riverside County, July 24, 1947, Aug. 24, 1947, 
MacSwain; 2, California (State label only); 29, Coachella, Riverside 
County, Apr. 28, 1927, May 9, 12, 17, 28, 1927, Wymore, May 22, 
1928, Van Dyke, May 13, 1917, Van Duzee; 2, Coachella Valley, 
Sept. 8, 1929, Comstock, Aug. 20, 1949, v. d. Bosch; 1, El Centro, 
Aug. 28, 1948, light trap; 8, Holtville, June 1936, Ross, August (no 
date given), Craig, June 28, 1936, Cazier, June 27, 1936, July 15, 
1936; 16 (includes allotype), Imperial County, June 1912, on Experi- 
ment Farm, Bridwell; 4, Imperial County, June 1925, Field, July 1, 
1925, Gehring; 2, Indio, July 17, 1923, Benedict, May 23, 1919, 
Slevin; 2, Meloland, June 4, 7, 1949, v. d. Bosch; 1, Needles, June 
12, 1940, Barr; 1, Neighbors, Apr. 27, 1930, Hornung; 1, Palo Alto, 
July 28, 1916; 1, Vallecito, San Diego County, Sept. 15, 1945, at light, 
Martin; 2, Warners, June 18, 1925. 

México: 3, 10 miles south of Catavina, Baja California, July 29, 
1938, Michelbacher and Ross. 

Paratypes in CAS, LAC, AMNH, CARN, INHS, CAL, OSU, CU, 
MICH, TEX, KAN, CDA, USNM, and the private collections of 
R. H. Arnett, D. K. Duncan, C. A. Frost, P. C. Grassman, Henry 
Howden, A. T. McClay, Gayle H. Nelson, F. H. Parker, Mark 
Robinson, and F. Werner. 

Remarks.—Males of B. imperialis vary from 11 to 19 mm. in length, 
and from 7 to 11 mm., in width. In lesser developed males the punc- 
tures become fewer and weaker over the posterior third inside the 
arcuate carina at back of the head, anteriorly they persist strongly; 


112 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


the clypeal horn decreases in height and its apex becomes narrow, 


sometimes rounded, but usually evidently truncate, and the anterior 


face vertical or sloping slightly backward; the median pronotal promi- 
nence is usually not constricted or narrowed basally. 

B. wmperialis is allied to B. serratus (LeConte), B. regalis Cartwright, 
and to B. peninsularis (Schaeffer), which I believe is a distinct species. 
Well-developed males of B. imperialis are separated from the other 


species by the anterior clypeal horn being relatively farther forward. 


from the lateral horns, the triangle formed by the tips of the three 
horns approaching an equilateral triangle much more closely than in 


the other species. The apex of the clypeal horn is much narrower — 
than in B. regalis, and usually truncate rather than rounded as in — 


serratus and peninsularis; in fully developed males the inner lateral 
carina is doubly angulate as in regalis but has the high middle peak 
bent inward, while in specimens of serratus and peninsularis this carina 
is not higher or angulate at middle. The color of serratus and penin- 
sularis is dark red-brown, of imperialis, a more yellow brown. 

In Bolbocerastes peninsularis, represented by 13 specimens, 3 of 
them cotypes, the male clypeal horn is rounded as in B. serratus but is 
much farther back relative to the lateral horns, which are nearly as 
high as the clypeal horn. The female is more heavily punctate, as in 
serratus, and the area back of the median horn is much more noticeably 
punctate than in wmperialis. 


Bolbocerastes imperialis kansanus, new subspecies 


Holotype male, length 17 mm., width 11 mm. Very similar to B. 
imperialis; moderately shining; darker in color;hairy beneath, Anterior 
face of the narrowly truncate clypeal horn vertical; lateral secondary 
horns more prominent; surface back of the clypeal horn with a notice- 
able swelling on each side and diagonally backward from the posterior 
ridges of the horn to a point even with the forward edge of the eye 
canthus, leaving between them a widening depressed area which 
merges with the general posterior level of the head; surface anterior 
to the arcuate ridge between the eyes more concave and much smoother 
than in typical imperialis; occiput smooth; eye canthus with a swelling 
diagonally inward from outer anterior angle. 

Pronotum more coarsely punctate throughout; lateral edges of the 
median prominence more nearly cariniform and not constricted 
basally; median peak of upper lateral carina not bent inward; the 
depression back of the anterior angles at bottom of the lateral sinuses 
deeper, more noticeable, and distinct; serrations of the lateral margin 
appearing as thickenings of a single sharp edge; otherwise similar to 
typical imperialis. 

Elytral striae deeper, nearly as in B. serratus (LeConte) ; the second 
stria basally indistinct over twice the length of the scutellum. 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 113 


Aedeagus as in wmperialis. 

Allotype female, length 16 mm., width 11 mm. Similar to typical 
B. imperialis except that punctures are coarser throughout; foveae 
are deeper; elytral striae are deeper; and the eye canthus has a convex 
swelling diagonally inward from the anterior angle. 

Type.—Holotype male, in Canadian Department of Agriculture 
collection. 

Type locality Rush County, Kan. 

Allotype and paratypes.—Allotype and 6 paratypes bear data as 
follows: Kansas: 4 (including allotype female), Rush County, Aug. 5, 
1920; 2, Ness County, Aug. 5, 1920; 1, Ness County, 2,260 feet, 
July 5, 1912, Williams. Paratypes vary from 13 to 17 mm. in length, 
and 8 to 11 mm. in width. 

Paratypes in CDA, KAN, and USNM. 

Remarks.—Bolbocerastes imperialis kansanus may be separated from 
B. serratus (LeConte) by the doubly angulate upper lateral thoracic 
carina in well-developed males, by the diagonal convexity of the eye 
canthus, by the lateral serrations of the pronotum appearing as 
thickenings of a single sharp margin, and by the second elytral stria 
all but disappearing at twice the length of the scutellum. In serratus 
the serrations of the pronotal margin arise between the two edges of 
the doubly carinate margin and the second elytral stria becomes 
indistinct opposite the apex of the scutellum. 


Bolbocerastes serratus (LeConte), new combination 
Fiaure 14, f; Puate 4 


Athyreus serratus LeConte, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, p. 80, 1854. 

Amechanus serratus Horn, Trans. Amer. Ent. Soc., vol. 3, p. 48, 1870. 

Bradycinetus serratus Horn, Proc. California Acad. Sci., ser. 2, vol. 4, p. 334, 1894. 

Athyreus (Bradicinetus) serratus Boucomont, in Wytsmann, Genera insectorum, 
fasc. 7, p. 8, 1902. 

Bradycinetus serratus Schaeffer, Trans. Amer. Ent. Soc., vol. 32, p. 251, 1906. 

Bradycinetulus serratus Cockerell, Ent. News, vol. 17, p. 242, 1906. 

Bolboceras (Amechamus) serratus Boucomont, Ann. Soc. Ent. France, vol. 79, 
p. 341, 1910. 


Bolbocerastes serratus (LeConte) and B. imperialis Cartwright, ex- 
cept for fully developed males, are frequently difficult to separate. 
The locality label will help in most cases since serratus is found east of 
New Mexico, while imperialis is found in California, Arizona, and 
rarely into west Texas. The maps (figs. 15 and 16) show the known 
distribution. In doubtful cases the male genitalia will separate the 
two (see fig. 14, f and g). In serratus, the basal side or floor of the 
cavity is formed by the two sides approaching each other with knife- 
edges, sharply angulate near the outside margin (in Bolbocerastes the 
aedeagus of the male is hooded apically by the two sides coming together 


114 , PROCEEDINGS OF THE NATIONAL MUSEUM | VOL. 103 


e Bolbocerastes imperialis Cartwright x Bradycinetulus rex Cartwright 


o Bradycinetulus fossatus (Haldeman) 


@ Bolborhombus schaefferi (Boucomont) © Bolbelasmus arcuatus (Bates) 


x Bolbocerastes peninsularis (Schoeffer) ~ 


Ficure 15.—Distribution of species of Bradycinetulus and close'y related species. 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 115 


© Bolbelasmus horni (Rivers) © Bradycinetulus ferrugineus (Palisot de Beauvois) 


® Bolbelasmus minor (Linel!) ® Bolbocerastes Imperialis kansanus Cartwright 


e Bolbocerastes regalis Cartwright © Bolborhombus angulus (Robinson) 


@ Bolbocerestes serratus (Leconte) x Bolborhombus parvulus Cartwright 


Ficure 16.—Distribution of species of Bradycinetulus.and closely related species. 


116 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


to form a deep cavity) ; in imperialis, these edges are rounded and lead 
straight in without a break or angle. 

Type.—In LeConte collection, MCZ. 

Type locality—Mexican Boundary, Laredo to Ringgold Barracks. 

Specimens examined.—181. 

Distribution —Kansas: Clark County. Oklahoma: Taloga, Chey- 
enne, *Alva, and *Payne County. Texas: Brownsville, *Beeville, 
Burnet County, Cypress Mills, Comal County, Cameron County, 
Caterina, Colorado City, Cotulla, Childress, Dimit County, Dallas, 
Dallas County, Edinburg, Eagle Pass, Fedor, Forestburg, Frio State 
Park, Goldthwaite, Hidalgo County, Kingsville, Laredo, Memphis, 
Menard, *Mineral Wells, New Braunsfels, Oakville, Plesanton, Reeves 
County, Round Mountain, Sabinal, Taloga, Uvalde, Weslaco, and 
Zavalla County. New Mexico: State label only (H. W. Wenzel col- 
lection). México: La Gloria, south of Monclova, Coahuila; and 
Rancho Preso, Nuevo Leén. 

Season.—April 28 to September 25, with largest numbers collected 
in May, June, and September. 


Bolbocerastes peninsularis (Schaeffer), new combination 


Figure 14, d; Puatr 4 


Bradycinetus serratus var. peninsularis Schaeffer, Trans. Amer. Ent. Soc., vol. 32, 
p. 252, 1906. 

Bolboceras serratus var. peninsularis Boucomont, in Schenkling, Coleopterorum 
catalogus, pt. 46, p. 13, 1912. 

All specimens of this species examined have been dull, smooth, and 
of a uniform very dark red-brown color. The male is quite distinctive 
(see pl. 4, lower right). In the female the moderately high cephalic 
horn is twice the width of the tip of the clypeal horn and the sec- 
ondary lateral horns or tubercles are usually more prominent than in 
B. imperialis. The carinae between these and the clypeal horn are 
strong and well marked. 

Type.—Lectotype male, present selection, USNM 42568. 

Type locality—Santa Rosa, Baja California, México. 

Specimens examined.—13. 

Distribution.—California: 2, State label only, CARN. México: 
Santa Rosa, San Felipe, La Paz, and San Ignacio, all in Baja Cali- 
fornia. 

Season.—July 27 to October 1. 


Bolborhombus, new genus 


Mandibles arcuate externally. Head, male and female, without 
median frontal horn. Pronotum serrate laterally; apex with a carina 
paralleling anterior margin, the interspace interrupted behind each 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 117 


eye by a distinct rather deep foveola; base margined. Scutellum tri- 
angular, with base straight, sides arcuate. Elytra not margined at 
base; seven discal striae, the first two interrupted by the scutellum, 
the second sometimes indistinctly forked opposite the apex of scutel- 
lum. Two prosternal spines in tandem behind anterior coxae, the 
posterior more or less hastate. Metasternal plate rhomboid in shape, 
the posterior angle acute and with adjacent edges cariniform, the 
lateral angles with adjacent edges rounded; the intercoxal lobe deeply 
concave and with strong cariniform lateral edges. 

Genotype.—Bradycinetus carinatus Schaeffer, 1906 = Bolboceras 
schaefferi Boucomont. 


Key to the species of Bolborhombus 


1. Clypeus of male carinately margined in front, the very wide clypeal horn 
arcuate anteriorly, a low sharp carina arcuate in opposite direction behind 
the clypeal horn, which is midway between this and the carinate anterior 
mmr deranie daknown so 2220 flo 3 2225 22k angulus (Robinson) 

Male clypeus not distinctly carinate along anterior edge but with parallel 
earinae extending back from anterior angles to the external angles of the tip 
Sy ana ENCE CD ToT sd are re eee he ee fn ts nt ee eee ee ne eee = 2 

2. Postapical carina of pronotum sharp and slightly undercut; male with short, 
distinct carina back of clypeal horn arcuate anteriorly, in same direction 
as clypeal horn; female smoother back of corresponding carina than in 
front of it; small shining species; 8to 10mm. (Baja California). 

parvulus new species 

Postapical carina of pronotum rounded; male without carina or with faint 
indication of two low obscure tubercles back of clypeal horn; female equally 
rough behind and in front of carina; usually dull or only moderately shining 
(excepting some very large specimens, which may represent a subspecies) ; 
muaperspecies. tito 1S mm- 2.42202 2 3 eo schaefferi (Boucomont) 


Bolborhombus angulus (Robinson), new combination 


FiaureE 14, b 


Bolboceras angulus Robinson, Trans. Amer. Ent. Soc., vol. 73, p. 170, 1947. 


Bolborhombus angulus (Robinson) is a small rufotestaceous species, 
7 to 11 mm. in length, readily recognized by the form of the head and 
clypeus. The wide straight anterior edge of the clypeus is strongly 
carinately margined. Behind this, from side to side and almost in 
line with the anterior edges of the ocular canthi, is a moderately high 
anteriorly arcuate ridge or horn, and still farther back a lower poste- 
riorly arcuate sharp carina. The median horn tends to become some- 
what nodose at the ends and is approximately midway between the 
posterior carina and the carinate anterior edge of the clypeus. In 
other characters angulus is very similar to B. parvulus and to small 
specimens of B. schaeffert (Boucomont). 

Type.—In Mark Robinson collection, Philadelphia, Pa. 


118 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Type locality —Dog Canyon, Brewster County, Tex. 

Specimens examined.—4, including the holotype. 

Distribution.—Texas: Dog Canyon, Glenn Spring, and Boquillas, 
all in Brewster County. Arizona: Miller Canyon, Huachuca 
Mountains. 

Season.—July 7 to 29. 


Bolborhombus parvulus, new species 


Fiaure 14, a 


Holotype male, length 9 mm., width 6mm. Shiny, rufotestaceous, 
densely hairy beneath. Clypeus extending obliquely upward and 
backward to top of clypeal horn; the anterior face practically flat, 
its surface closely, very coarsely punctate, with the forward edge 
convex and deeply reticulate-punctate, the lateral edges parallel and 
sharply cariniform; upper edge of the horn shallowly emarginate, the 
external angles only slightly tuberculiform, from them a moderately 
high, sharp carina curving back each side and ending somewhat 
abruptly close to the innermost edge of the eye; surface of head back 
of horn uneven, with close, mixed fine and coarse punctures and with a 
short, elevated, transverse, anteriorly arcuate carina midway between 
the similarly punctate, sharply edged, concave, depressed ocular canthi; 
the carina about one-third as long as the width of the clypeal horn; 
the punctures back of the carina not quite as close as in front; the 
occiput smooth, closely, minutely punctate. 

Pronotum with three rather deep foveae close behind the finely 
margined anterior edge, one median, the others opposite inner eye 
margin on each side; a sharp postapical carina, slightly undercut on 
posterior side, starting on each side of the middle fovea and passing 
close behind the lateral foveae toward but not quite reaching the 
serrate side margin at about anterior third; a second short, similar 
carina on each side straight back from near the end of the postapical 
carina to the inner anterior edge of a strong lateral fovea; a shallow, 
coarsely punctate groove half way across the pronotum upward and 
backward from each of the three anterior foveae; a row of close coarse 
punctures in the lateral and basal marginal line, a few m the basal 
half of median line, and scattered coarse punctures on both sides of 
the lateral carina and continued upward to opposite the humerus, with 
mixed fine and minute punctation throughout; anterior angles slightly 
more than a right angle; posterior angles obtusely rounded; base 
distinctly margined. Scutellum with mixed, very fine and minute 
punctation. 

Elytra not margined basally; striae weakly impressed; strial punc- 
tures fine; first two striae interrupted by the scutellum; intervals 
weakly convex; the surface minutely punctate, very finely alutaceous 
under moderately high magnification. 


BOLBOCERATINI OF THE UNITED STATES—CARTWRIGHT 119 


Metasternal plate twice as wide as the lobe between the middle 
coxae, roughly punctate, and hairy along posterior edges. Prosternal 
spine 4-cornered, with the anterior angle curving down to a sharp 
point between the coxae and with the lateral angles on a transverse 
level and the posterior curving downward to form a sharp spine. 

Allotype female, length 9 mm., width 6mm. Apparently identical 
with the male in every way except in characters of the head. Clypeus 
rising steeply at somewhat less than a right angle to a sharply carinate 
edge seven-eighths as wide and parallel to the turned-under anterior 
margin; the flat face enclosed being a little less than one-third as high 
as long, then dropping sharply and gradually up again with the 
lateral edges slightly diverging to the top of the very wide median 
horn, which is widely and deeply emarginate at middle, leaving the 
two ends, or external angles, as rounded, thickened, backward-sloping 
prominences; back of the emargination, on the same level, and at a 
point midway between the ocular canthi, is a very short arcuate 
carina or elongate tubercle, behind which the general level of the head 
is noticeably lower; surface in front of the horn deeply, coarsely 
punctate-reticulate, between the horn and the tubercle roughly 
coarsely punctate, and behind the tubercle much smoother, with 
dispersed, mixed fine and shallow, coarse punctures; occiput smooth, 
practically impunctate. 

Type.—In California Academy of Sciences collection. 

Type locality —Triunfo, Baja California, México. Holotype male 
collected Aug. 7, 1938, Michelbacher and Ross. 

Allotype and paratypes.—Allotype female, CAS, and paratype male, 
USNM 61656, San Venancio, Baja California, México, Oct. 8, 1941, 
Ross and Bohart; paratype female, USNM 61656, Santa Rosa, Baja 
California, México, Charles Schaeffer collection. 

Remarks.—Bolborhombus parvulus is very near B. angulus (Robin- 
son) and is about the same size. The four specimens vary in length 
from 8% mm. for the smaller male to 10% mm. for the larger female. 
The anterior edge of the clypeus, described as margined in angulus, is 
almost exactly the same as in the female of parvulus. 


Bolborhombus schaefferi (Boucomont), new combination 
FicureE 14, ¢ 


Bradycinetus carinatus Schaeffer, Trans. Amer. Ent. Soc., vol. 32, p. 251, 1906. 
Bolboceras schaefferi Boucomont, Ann. Soc. Ent. France, vol. 79, p. 347, 1910. 


Bolborhombus schaefferi (Boucomont) averages much larger than 
B. parvulus and B. angulus, measuring 11 to 18 mm. in length, but it 
is very similar in all characters except those of the head, clypeus, and 
aedeagus. Males show considerable variation in the clypeal horn. 


120 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


In smaller specimens the clypeus slopes upward and backward from — 


the wide straight anterior edge to the narrower but moderately wide 
and somewhat anteriorly arcuate apex of the horn, the external 
angles of which are thickened to slightly nodose. In large specimens 
the external angles of the horn become strongly elevated tubercles, in 
some individuals wide apart, in others relatively close together. The 
head back of the horn is densely, quite coarsely punctate, occasionally 
with a short, obscure, anteriorly arcuate carina or very low binodose 
tubercle. In the female the anterior edge of the clypeus is nearly 
vertical to the sharply carinate margin. Well back and nearly in line 
with the anterior edges of the ocular canthi are two widely separated, 
conical tubercles with a fine anteriorly angulate carina between them 
and a low median swelling or tubercle close behind them. The 
tubercles are usually strongly developed but occasionally in small 
specimens may appear only as slight thickenings of the ends of the 
angulate carina. 

Type.—Lectotype male, present selection, USNM 42569. 

Type locality —Palmerlee, Cochise County, Ariz. 

Specimens examined.—104. 

Distribution—Arizona: Cave Creek and Pinery Canyon (Chira- 
cahua Mountains, Cochise County), Dewey, Douglas, Globe, Hua- 
chuca Mountains, Fort Huachuca, Kits Peak (Rincon, Baboquivari 
Mountains), Nogales, *Oracle, Palmerlee, Patagonia, Prescott, 
Reddington, Madera Canyon (Santa Rita Mountains), and Tucson. 
Texas: *Alpine, El Paso, *Fort Davis, Kingsville, and Round Moun- 
tain. México: Alamos (Sonora), Venedio (Sinaloa), Oaxaca (Oaxaca), 
Santa Rosa (Baja California), Naco (Sonora), Tlahualilo, and Rio 
Mayo. 

Season.—July 1 to October 10, with largest numbers in July and 
August. 


U. $. GOVERNMENT PRINTING OFFICE: 1953 


U. S. NATIONAL MUSEUM 


PROCEEDINGS, VOL. 103, PLATE 3 


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Front and dorsal views of three species of Bradycinetulus: Left, B. fossatus (Haldeman); 
middle, B. rex, new species; right, B. ferrugineus (Palisot de Beauvois). 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 4 


Differences in the clypeal horn in Bolbocerastes: Upper left, B. serratus (LeConte); upper 

: penyelt : Nees ; eee 
right, B. regalis, new species; lower left, B. imperialis, new species; lower right, B. 
peninsularis (Schaeffer). 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1953 No. 3319 


THE CHRYSOMELID BEETLES OF THE GENUS 
STRABALA CHEVROLAT 


By Doris Hotmes BLAKE 


Introduction 


The first species of this chrysomelid genus was described by Illiger ! 
in 1807 as Haltica rufa from Pennsylvania, Melsheimer collector. 
A year later, Olivier? described <Altica ferruginea from Santo Do- 
mingo, Palisot de Beauvois collector, and Altica scutellaris from 
southern France. Of this latter locality von Harold * said that there 
was no Altica colored anything like this in southern France and that 
the locality was evidently an error. In 1837, in the Dejean catalogue, 
Chevrolat* listed under the new generic name Strabala (crpaBadra, 
having a form thick and contracted) six specific names of Dejean 
without descriptions or references. Four of them are pure nomina 
nuda; for the two others, S. scutellata Dejean and S. dominicensis 
Dejean, the names (Altica) scutellaris? Olivier and (A.) ferruginea? 
Olivier, respectively, are given as doubtful synonyms. The generic 
name is consequently not published here. Chevrolat changed the 
locality for scutellata to America Borealis. Of these six names only 
one has since been used; S. intermedia, whose locality in this catalogue 
was given as Cuba, was adopted by Jacquelin Du Val °* in 1857 for a 

1 Tiliger, Mag. Insekt., vol. 6, p. 152, 1807. 

2 Olivier, Entomologie, . . . vol. 6, pp. 697, 699, 1808. 

4 von Harold, Coleopterologische Hefte, vol. 14, p. 20, 1875. 

4 Ohevrolat, in Dejean, Catalogue de la collection de Coléoptéres . . . ed. 3, p. 413, 1837. 


* Jacquelin Du Val, in Ramé6n de Ja Sagra, Historia . . . de la Isla de Cuba (Spanish ed.), vol. 7, p. 129, 
1857. 


232989—53 121 


122 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Cuban species. In d’Orbigny’s Dictionnaire, Chevrolat® in 1848 
listed scutellaris and ferruginea under Strabala without the question 
marks, as well as the four nomina nuda he had given in 1837, thus 
establishing the genus. 

In 1868 Suffrian” described Haltica ambulans from Cuba which he 
said belonged to Chevrolat’s genus Strabala, a genus that Erichson ® 
had earlier merged with Lactica. Suffrian was not convinced, however, 
that these specimens having so poorly developed a basal sulcus on the 
prothorax really belonged to the genus Lactica. He stated that 
although there was an ill-defined basal sulcus, the beetles were smaller 
than most species of Altica and proportionately broader, and that the 
color was not blue but a deep reddish brown. This Cuban species that 
he had did not entirely correspond in coloring to Olivier’s description 
of scutellaris or to Jacquelin Du Val’s intermedia, both of which were 
paler beneath than the dark colored undersurface of his beetles. 

Boheman ® described under the genus Strabala, two species, nigriceps 
from Buenos Aires and languida from Java, both of which have been 
referred to Lactica. Although I have not examined the types, the 
description of nigriceps as having the anterior angles of the prothorax 
truncate seems to exclude it from Strabala, and the description of 
languida is quite different from that of any of the group. Fall,!® in 
describing Altica testacea from Texas, compared it with rufa, noting 
that ‘‘rufa is much broader and less convex and has black legs and 
antennae.” Fall’s species, also, does not belong in Strabala. It differs 
in general shape, the prothorax has quite different anterior angles, 
and the scutellum is small and inconspicuous. 

Jacoby,!! who had put scutellaris under Lactica, in the Supplement 
says of Horn, ‘Dr. Horn remarks that Lactica scutellaris would be 
better placed in the genus Haltica and that it is congeneric (and prob- 
ably also specifically identical) with Haltica rufa Illiger.” But 
Jacoby ‘‘could not agree with this opinion” and kept scutellaris under 
Lactica. 

The British Museum (Natural History) specimens, therefore, are 
under Lactica, and in the U. S. National Museum, following Horn, 
they are under Altica. Henshaw” lists rufa under Disonycha. 
Gemminger and Harold ® list rufa under Disonycha and _ scutellaris 
under Lactica. Heikertinger,' in the Junk catalogue, nicely balances 


8 Chevrolat, in d’Orbigny, Dictionnaire universe! d’histoire naturelle, vol. 12, p. 52, 1848. 

1 Suffrian, Arch. Naturg., vol. 34, p. 182, 1868. 

§ Erichson, Arch. Naturg., vol. 13, p. 178, 1847. 

® Boheman, Kongliga Svenska Fregatten Hugenies Resa. . ., vol. 2, Zoologi, pt. 1, Insecta, pp. 189, 190, 
1859. 

10 Fall, Trans. Amer. Ent. Soc., vol. 36, p. 157, 1910. 

11 Jacoby, Biologia Centrali-Americana, Coleoptera, vol. 6, pt. 1, p. 278, 1884; Supplement, p. 259, 1891. 

12 Henshaw, List of the Coleoptera of America, north of Mexico, p. 112, 1885. 

13 Gemminger and von Harold, Catalogus coleopterorum .. ., vol. 12, p. 3497, 1876. 

14 Heikertinger, Ooleopterorum catalogus, pars 166, pp. 241, 259, 1939. 


CHRYSOMELID GENUS STRABALA CHEVROLAT—BLAKE 123 


the dispute by synonymizing scutellaris and ambulans with rufa and 
placing them under Haltica and at the same time placing ferruginea 
and intermedia as separate species under Lactica. Mr. G. E. Bryant 
pointed out to me the specimens labeled Lactica scutellaris in the 
British Museum and said, ‘‘What are these doing here?” I agreed 
that it was high time to restore the original Chevrolat generic name 
Strabala for this group of very similarly colored species that no one 
has known exactly what to do with. On my return to America, after 
a considerable search for the specimens of this species that had long 
been removed from the regular collection, I found that H. S. Barber 
had also labeled them Strabala. 

The species of this genus differ in color among themselves little more 
than do those of the genus Altica, but instead of being metallic blue 
they are a deep reddish or orange brown with dark or partly dark legs, 
dark scutellum (usually), dark antennae, and more or less dark under- 
surface. Like Altica they have no spots, vittae, or other markings; 
and the variation in the dark coloration is slight. In practically all 
the North American specimens from Massachusetts to Panama, the 
amount of dark coloration is very much the same, that is, the under- 
surface and femora are only partly dark. Hence the tendency has 
been to follow Horn,” who wrote that ‘“Haltica rufa occurs from 
Massachusetts to Illinois, Florida and Texas, extending through 
Mexico to South America.” There are, however, slight differences in 
the extent of the dark coloration, in the punctation, and in the shape 
of the aedeagus. In Eastern United States from New Hampshire to 
the Gulf and west to the Mississippi, Louisiana, and northern Texas, 
the species known as rufa Illiger presents little variation. It is a con- 
spicuously punctate, deep reddish brown species. In Florida occurs a 
race that, in dried specimens, at least, is always a pale yellow-brown, 
in contrast to the reddish color of rufa. It has also a distinctly aluta- 
ceous surface and is less distinctly punctate than rufa. In the speci- 
mens from Brownsville, Texas, to Panama, which are shining and 
almost impunctate, in our limited collections are two groups with 
entirely differently shaped aedeagi—those that are narrowed at the 
tip and those with a wide, rounded tip. From the external appearance 
I cannot distinguish the beetles at all. H.S. Barber has labeled one 
lot, with round-tipped aedeagi, as a new species and the other as 
scutellaris. It seems to me that the name scutellaris cannot be applied 
to any species of Strabala with any certainty. To begin with, Olivier’s 
locality of southern France, apparently erroneous, leaves us without 
any type locality in the western hemisphere. Species corresponding in 
coloration to Olivier’s description occur in Cuba and Central America 
as well as North America. Therefore I propose to drop the name 


Horn, Trans. Am. Ent. Soc., vol. 16, p. 232, 1889. 


124 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


scutellaris entirely, as retaining it would only promote confusion. 
Among the specimens from Central America with the aedeagus nar- 
rowed at the tip there is a wide variation in the acuteness and in the 
width of the tip. I think there are probably a number of species, but 
since the coloring is so nearly uniform and the variation in the 
aedeagi only a matter of degree, I hesitate to give them specific names 
and am calling them only subspecies. 

In the Bowditch collection at the Museum of Comparative Zoology 
(MCZ) are seven specimens from South America (two from Venezuela 
and the rest from Colombia) that are uniformly somewhat larger than 
the others of the genus and, unlike the North and Central American 
specimens, are dark beneath, as in the majority of the West Indian 
specimens. In the British Museum is a series from Trinidad, col- 
lected by G. E. Bryant, also dark beneath but smaller, paler, and with 
a more rounded tip to the aedeagus. 

In the West Indies the prevailing color of the legs and undersurface 
of the specimens is entirely dark, but in one species, at least, the pos- 
terior femora are partly reddish, as in the continental species. In 
another species from Hispaniola, the scutellum is not black but 
reddish like the elytra, the only one so colored that I have examined. 
Jacoby wrote of specimens with a red scutellum from Juquila, México, 
of which he had three. One specimen from Juquila in his collection 
is a yellow Lactica with a pronounced basal sulcus on the prothorax, 
typical of Lactica, and a pale scutellum. This may not be the one to 
which he referred. These slight differences in coloration of the speci- 
mens from the different islands of the West Indies are accompanied 
by a slightly different aedeagus. Still, as in the case of the Central 
American specimens, I hesitate to give them specific names and am 
calling them only subspecies. Much more work should be done on 
the group and many more specimens should be studied. Here I 
attempt to assemble the species that in the catalogues are divided 
among two or three genera and to call attention to their points of 
difference. 

The group is closer to Disonycha than to either Altica or Lactica. 
The head, while similar to Disonycha, has the circle of punctures 
nearer to the tubercles and the carina shorter and broader than in 
most species of Disonycha. The antennal joints are similar in being 
robust and rather short, but differ in having the third joint a little 
longer than the fourth, instead of the fourth longer than the third. 
The prothorax is much like that of Disonycha except that below the 
apical angle there is a tiny V-shaped notch which does not occur in 
Disonycha. The basal depression over the scutellum, often found in 
Disonycha too, is longer and more pronounced in Strabala. The 
elytra are distinctive in being more convex and oval than in Lactica, 
Altica, or Disonycha. 


CHRYSOMELID GENUS STRABALA CHEVROLAT—BLAKE 125 


g-S.ambulans (Sef friar) 4.S.ambulans ssp jarnalcensis i. S. Ferrugtnea Olivier) j-S.ambulane ssp. puertoriceasis 


k. Scolumpiana LS.intermedia Jacq.duVab m S.frinitalis 


Ficure 17.—Species of Strabala Chevrolat: a, rufa (Illiger); b, rufa floridana, new subspecies; 
€,acuminata, new species; d, rotunda, new species; ¢, acuminata teapensis, new sub- 
species f, acuminata costaricensis, new subspecies; g, ambulans (Suffrian); h, ambulans 
jamaicensis, new subspecies; i, ferruginea (Olivier); 7, ambulans puertoricensis, new 
subspecies; k, columbiana, new species; 1, intermedia Jacquelin Du Val; m, trinitatis, 
new species. 


126 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Genus Strabala Chevrolat 


Description.—Between 3.5 and 6 mm. in length, oval, deep yellowish 
or orange brown or reddish brown, with antennae dark, legs and under- 
surface more or less dark, and scutellum usually conspicuously dark; 
moderately shining, most species very minutely and confusedly 
punctate, only the North American species, S. rufa (Illiger), very 
distinctly punctate. 

Head, except for mouth parts that are usually a little darker, deep 
yellowish or reddish, eyes entire and widely separated, the interocular 
space being more than half the width of the head; occiput smoothly 
rounded down to tubercles, with scattered and not very dense or coarse 
punctures; at the base of occiput, not visible except when the head is 
a little protruded, a groove running from behind one eye to the other; 
on either side above the frontal tubercles and near the eye a depression 
consisting of a large fovea, or more frequently a circle of punctures; 
frontal tubercles plainly marked, a broadly rounded and somewhat 
produced area between antennal sockets that extends and widens down 
the front nearly to the labrum, area under sockets a little depressed 
and on sides below eyes the cheeks rounded out and densely punctate. 
Antennae rather stout, not quite reaching the middle of the elytra, 
first two joints swollen, usually paler, remainder piceous, third joint 
a little longer than fourth, fourth and fifth about equal. Prothorax 
not quite twice as wide as long, with rounded sides and narrow margin, 
the apical angle resembling that of Disonycha except that it is sepa- 
rated from the rest of the margin by a distinct though tiny V-shaped 
indentation, basal angle with a small tooth and obliquely cut, disk 
smooth and rather flat, with more or less of a depression along base 
and not distinctly limited at the ends as in Lactica. Scutellum large, 
usually conspicuously dark, contrasting with the reddish elytra. 
Elytra broader than prothorax, moderately convex, with small 
humeri and narrow margin, not depressed; surface usually smooth, not 
distinctly punctate except in S. rufa, entirely yellowish or reddish 
brown. Epipleura broad, gradually diminishing and not quite reach- 
ing the apex. Body beneath variable in color, the prosternum red- 
dish, the breast and abdomen and legs in West Indian species usually 
but not always dark, in North American species the breast dark on 
the sides, the femora pale except in apical half or sometimes entirely 
pale, the tibiae and tarsi dark, the sides and apex of the abdomen 
sometimes also dark; in Trinidad and South American species, the 
coloring beneath dark as in the West Indian specimens. Anterior 
coxal cavities open, front coxae narrowly separated, legs rather short, 
hind femora moderately thickened, the tibiae as in Disonycha with a 
slight ridge, a spur at the end of posterior tibiae, claws appendiculate. 


CHRYSOMELID GENUS STRABALA CHEVROLAT—BLAKE 127 


Aedeagus a simple, curved structure with the tip varying in shape 
from acute to truncate and broadly rounded. 
Genotype.—Strabala ferruginea (Olivier), present designation. 


Key to the species of Strabala 


i -peutellum. reddish (Hispaniola) -:-~- 222222 2. 2-2 ferruginea (Olivier) 
Scucclum dark brown) OF piceoUShis heey Bre val wees eee 2 

2. Species from the West Indies exclusive of Trinidad______.._..-------_-- 3 
Beeies. roms Nomn AmMeniCe 420% AN 5 
Species from South America and Trinidad__.__..._-___..-~+---+------- 9 

3. Body beneath more or less ferrugineous, hind femora ferrugineous at base, 
Bare niraApex (GUDR)s 2 eh 8 he STN 5 intermedia Jacquelin Du Val 
Body beneath except prosternum more or less dark, hind femora entirely 
penne wwe ore th SER otis el eho drupe bya. py pepe 4 


4, Aedeagus tapering gradually to a rounded tip (Cuba)-.---ambulans (Suffrian) 
Aedeagus tapering gradually to a more acute but still rounded tip (Jamaica). 
ambulans jamaicensis, new subspecies 

Aedeagus distinctly truncate at tip (Puerto Rico). 
ambulans puertoricensis, new subspecies 
5. Elytra punctate or alutaceous or both (North of México)_-___---------- 6 
Elytra not distinctly punctate or alutaceous (Texas and southward) ----_- 7 
6. Elytra deep reddish brown, distinctly punctate (New Hampshire south and 
westward to the Mississippi, Louisiana, Texas, and along the Gulf to Florida). 
rufa (Illiger) 
Elytra (in dried specimens) yellow, less distinctly punctate, alutaceous 


(TDG TUG UE) ey Rat fo rn on ty leds rufa floridama, new subspecies 

7. Aedeagus well rounded at tip and with a slight constriction behind the tip 
GViexico.to)Panams) ites yeeie em e reat) aries whee rotunda, new species 
Aedeagus gradually narrowing towards the tip, not constricted behind the 
£1 a a es ee ER eA en ey ete ae eee er eee 8 


8. Aedeagus narrowed to a rounded tip, hind femora dark at apex (Texas). 
acuminata, new species 
Aedeagus narrowed to a truncate tip, hind femora entirely reddish brown 
PexICg) 2 seo oe Joe acuminata teapensis, new subspecies 
Aedeagus narrowed to a broader truncate tip, hind femora varying from 

entirely reddish to reddish with a dark apex (Guatemala, Costa Rica). 
acuminata costaricensis, new subspecies 

9. Large, 5 to 6 mm., aedeagus with an acute tip (Colombia, Venezuela). 

colombiana, new species 

Smaller, 4 to 5 mm., aedeagus with a somewhat truncate tip (Trinidad). 
trinitatis, new species 


Strabala rufa (liger) 
Figure 17, a 
Haltica rufa Illiger, Mag. Insekt., vol. 6, p. 152, 1807. 


Illiger’s description of the thickly punctate elytra distinguishes 
this species from the others of the genus. The coloration is usually 


128 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


deep reddish brown above, with entirely dark antennae, dark scutel- 
lum, often entirely dark anterior legs, dark apical end of the posterior 
femora, dark breast, and sometimes also dark abdomen. Throughout 
its range over the country east of the Mississippi and in Louisiana and 
Texas, there seems to be very little variation except for the paler, 
less distinctly punctate race that occurs in Florida. The type locality 
is Pennsylvania, collected by Melsheimer. 

Distribution —New Hampshire: Hampton, S. A. Shaw. Massachu- 
setts: Holyoke, Chicopee, F. Knab. Connecticut: Hartford, F. Knab. 
Rhode Island: Watch Hill, W. Robinson. New York: West Point, 
W. Robinson. New Jersey: Boynton. Illinois: C. Thomas. Mich- 
igan: Detroit, Hubbard and Schwarz. Maryland. Washington, 
D. C. Virginia: Fredericksburg, Clifton, J. C. Bridwell; Nelson 
County, W. Robinson; Stone Creek, Lee County, Hubbard and 
Schwarz; Vienna, J. C. Bridwell; West Falls Church, E. A. Chapin. 
West Virginia: Aurora. Kentucky: H.Soltau. Tennessee: Memphis, 
H. Soltau. North Carolina: Graybeard Mountain. South Carolina: 
Florence. Georgia: Savannah. Alabama: Coleta, H. M. Smith. 
Mississippi: Greenwood, T. H. Parks. Louisiana: Baton Rouge, 
T. H. Jones; Port Allen, J. L. Lauderdale; New Orleans, H. Soltau. 
Texas: Liberty, R. A. Vickery. Two specimens, both females, 
USNM (Brooklyn Museum collection), labeled simply “Mexico,” are 
apparently this species. 


Strabala rufa floridana, new subspecies 
Ficure 17, 6 


This race differs from the reddish brown one found elsewhere in 
eastern United States by being (in dried specimens) pale yellow-brown 
with the dark markings as in typical rufa. The elytra are less dis- 
tinctly punctate than in rufa and distinctly alutaceous. 

Type—Holotype male and 7 paratypes, USNM 61202, labeled 
simply “Fla.,”’ collector F. Knab. 

Distribution.—Florida: Lake Ashby, Capron, Indian River, all 
collected by Hubbard and Schwarz; Lake Wales, Lakeland, E. M. 
Craighead; Jacksonville, Ashmead; Sanford, Van Duzee; St. Peters- 
burg (on sweet corn); Ft. Myers (on sweet and white potato); Oneco 
(on spinach); Homestead (on string beans); West Palm Beach (on 
sweet potato and lima beans). 


Strabala acuminata, new species 
Figure 17, c 


From 4.5 to 5.5 mm. in length, oval, shining, orange-brown, with 
dark antennae, dark scutellum, darkened sides to breast, and dark 
anterior legs and apical part of posterior femora, tibiae, and tarsi. 


CHRYSOMELID GENUS STRABALA CHEVROLAT—BLAKE 129 


Head shiny, only a few scattered punctures over occiput besides 
the fovea or circle of depressed punctures near the eye; mouth parts 
frequently edged with deeper brown. Antennae deep brown, with 
the three basal joints frequently paler. Prothorax shiny, almost 
impunctate, only under high magnification are very fine punctures 
visible; basal sulcus usually not very long or pronounced, but somewhat 
variable in different specimens, as is usual in the genus. Scutellum 
dark. Elytra mirror smooth, shining, impunctate, sometimes very 
indistinctly alutaceous. Body beneath with sides of breast, anterior 
legs, apical part of posterior femora, tibiae, and tarsi dark. Length 
4.4 to 5.5 mm.; width 2. 4 to 3 mm. 

Type.—Holotype male and 2 paratypes, USNM 61203 (Brooklyn 
Museum collection), collected at the Esperanza Ranch, Brownsville, 
Texas, August 18 (no year given). 

Other localities —Texas: Brownsville, R. A. Vickery; another speci- 
men, from same locality (on string bean leaves); San Benito (on 
bean foliage); Hidalgo (on black-eyed-pea leaf). 

Remarks.—The shiny, almost impunctate surface with little evidence 
of alutaceous dullness at once differentiates this from Strabala rufa 
(Illiger). There are no records of it farther north than the Brownsville 
region. 


Strabala acuminata teapensis, new subspecies 
FicurE 17, e 


Lactica scutellaris Olivier, Jacoby (part), Biologia Centrali-Americana, vol. 6, 
pt. 1, p. 273, 1884. 

This differs from the Brownsville, Texas, race in having less dark 
coloring, the femora all being deep reddish, the tibiae and tarsi only 
dark, and the sides of the breast not so dark. The upper surface is 
a shining deep reddish brown. The aedeagus has a somewhat trun- 
cate tip. 

Type.—Holotype male, in the British Museum (Natural History), 
from the Biologia Centrali-Americana material, labeled by Jacoby 
Lactica scutellatis Olivier, from Teapa, Tabasco, México, collected in 
March by H. H. Smith; 1 paratype in U. S. National Museum, 
USNM 61204; and 2 paratypes in Museum of Comparative Zoology 
from same series. 


Strabala acuminata costaricensis, new subspecies 


Figure 17, f 


Lactica scutellaris Olivier, Jacoby (part), Biologia Centrali-Americana, vol. 6, 
pt. 1, p. 273, 1884. 


This differs from the Brownsville, Texas, and Teapa, México, races 
by having a still wider truncate tip to the aedeagus. In coloring, it 


130 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


is somewhat deeper red than the Brownsville specimens and differs 
from the Teapa specimens in having the anterior femora and apical 
part of the posterior femora as well as the sides of the breast dark. 

Type.—Holotype male and 1 paratype, USNM 61205, collected at 
Vara Blanca, Costa Rica, 1,700 meters, July 7, 1928, by F. Never- 
mann; 1 paratype in British Museum (Natural History). 

Other localities —Costa Rica: San José, 1,000—2,000 meters, Aug. 25, 
1928, F. Nevermann. Guatemala: Yepocapa, August 1948, H. T. 
Dalmat; Cobén, Alta Vera Paz, May 1926, J. M. Aldrich; Purula, 
Champion 


Sirabala rotunda, new species 


Ficure 17, d 


Lactica scutellaris Olivier, Jacoby (part), Biologia Centrali-Americana, vol. 6, 
pt. 1, p. 273, 1884. 


From 4 to 5.5 mm. in length, oval, shining although faintly alu- 
taceous, very finely punctate, yellowish or reddish brown with dark 
antennae, dark apex to femora, dark tibiae and tarsi, and the breast 
and abdomen often darkened on the sides, varying to entirely dark 
(rare). 

Head reddish, with the mouth parts usually deeper brown; finely 
punctate over occiput, with the usual fovea or circle of depressed 
punctures. Prothorax shiny and very finely punctate with a sulcus 
along the base well developed. Scutellum dark. Elytra faintly 
alutaceous and finely punctate. Body beneath varying in color 
from having the sides of breast and abdomen deeper brown to being 
entirely dark, the latter rare; femora with apical half dark, tibiae and 
tarsi dark. Length 4 to 5.4 mm.; width 2.6 to 3 mm. 

Type.—Holotype male, USNM 61206, and 3 paratypes in British 
Museum (Natural History) collected at Volcén de Chiriqui, Panamé, 
by Champion. 

Other localities —Panam&: Panamd, Aug. 13, 1946, by N. L. H. 
Krauss; Almirante, Sept. 4, 1938, H. Dybas; Bella Vista, Aug. 7, 1924, 
N. Banks; Mindi Dairy (on para grass), Mindi, Canal Zone, July 
1918, H. F. Dietz and J. Zetek; Plantation Borracho, Canal Zone 
(swept from grass under papaya trees), July 1918, H. F. Dietz and 
J. Zetek; Bugaba, Champion (in British Museum). Guatemala: 
Trece Aguas, Alta Vera Paz, May (date not given), H. S. Barber and 
E. A. Schwarz. México: In bananas from México taken at Mobile, 
Alabama, July 19, 1929; with orchids from Guerrero (State), taken at 
Laredo, Texas, Feb. 28, 1940; with orchids from Tamazunchale, 
San Luis Potosi, taken at Laredo, Texas, Feb. 28, 1940; with tomatoes 
from E] Mante, Tamaulipas, taken at El Paso, Aug. 1, 1947; Colima, 


CHRYSOMELID GENUS STRABALA CHEVROLAT—BLAKE 131 


Hoge and Conradt; Jalapa, Hége; on green peppers from Manuel, 
Tamaulipas, taken at Laredo, Texas, Jan. 7, 1935. 

Remarks.—H. S. Barber has attached a specific name from the 
Mexican localities to this species, but since the range is as far south 
as Panamé, his name would be somewhat misleading. The rounding 
of the tip of the aedeagus varies considerably but the slight con- 
striction behind the tip is always present, and sets the species apart 
from the rest. The only other species with a similar aedeagus tip 
in the group is Strabala intermedia Jacquelin Du Val, from Cuba. 
In all, 15 specimens of S. rotunda have been taken on orchids 
from México at the port of entry to this country. 


Strabala ambulans (Suffrian) 


FicureE 17, g 
Haltica ambulans Suffrian, Arch. Naturg., vol. 34, p. 182, 1868. 


Specimens from Cuba agree with Suffrian’s detailed description of 
his Cuban H. ambulans. The upper surface is faintly alutaceous and 
finely punctate, in color a deep reddish brown with dark scutellum 
and dark legs, breast, and abdomen. 

Distribution Cuba: Cayamas, E. A. Schwarz; Central Jaron, 
L. C. Scaramuzza; Baragua, L. C. Scaramuzza, L. D. Christenson 
(MCZ); upper Yara Valley, Oct. 18, 1928, L. C. Scaramuzza (MCZ) ; 
Soledad, Cienfuegos, Nov. 7, 1926, P. J. Darlington (MCZ); Aguada 
(MCZ); Mountains north of Imias, eastern Oriente Province, July 
25-28, 3-4,000 feet, P. J. Darlington (MCZ). 


Strabale ambulans jamaicensis, new subspecies 


FicureE 17, h 


Between 3.5 and 4.5 mm. in length, oval, faintly shining, indis- 
tinctly alutaceous, very finely punctate, reddish brown, the mouth 
parts, antennae, legs, breast, and abdomen dark. 

Head with scattered punctures, the usual fovea or circle of punc- 
tures between eye and frontal tubercle on each side. Antennae with 
the basal joints a little paler. Prothorax having a distinct sulcus 
along the base. Scutellum dark. Elytra very faintly punctate. 
Body beneath with breast, abdomen, and legs dark. Length 3.5 to 
4.5 mm.; width 2 to 2.3 mm. 

Type—Holotype male and 10 paratypes, USNM 61207, collected 
at Manchioneal, Jamaica, Jan. 30, 1939, by E. A. Chapin and R. E. 
Blackwelder. 

Other localities —Jamaica: Roaring River Falls, St. Ann’s Parish, 
Sept. 13, 1917, H. Morrison; Fern Gully, St. Ann’s Parish, Sept. 14, 


132 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


1917, H. Morrison; Mandeville, Van Duzee, A. E. Wight (MCZ); 
Montego Bay, Mar. 2, 1911; Balaclava (MCOZ). 

Remarks.—This subspecies differs only slightly from the Cuban race 
in having a more acute though still rounded tip to the aedeagus. 


Strabala ambulans puertoricensis, new subspecies 
Figure 17, 7 


Between 4.5 and 5 mm. in length, oval, shiny, elytra not alutaceous, 
very indistinctly punctate, reddish brown, antennae, mouth parts, 
legs, breast and abdomen dark. 

Head with scattered punctures and a fovea or circle of punctures 
near the eyes. Antennae with paler basal jomts. Prothorax not at 
all clearly depressed along the base. Scutellum dark. Elytra 
quite shiny, not alutaceous, very finely and indistinctly punctate. 
Body beneath dark except the prosternum. Length 4.5 to 5 mm.; 
width 2.5 to 3 mm. 

Type.—Holotype male and 1 paratype, USNM 61208; 1 paratype 
in Museum of Comparative Zoology, taken at Adjuntas, Puerto Rico, 
Nov. 1, 1932, by R. G. Oakley. 

Other localities —Puerto Rico: Lares, September 1921, G. N. Wol- 
cott; Arecibo, June 17, 1932 (on eggplant leaf), Anderson and Mills; 
Mayagiiez, January 1899, A. Busck; Villalba, June 18, 1934, C. M. 
Matos (MCZ); Mayagiiez, December 1932, V. Alexandrina (MCZ). 

Remarks.—The shiny, not alutaceous elytra, the larger size, and the 
broad, almost truncate tip of the aedeagus are unlike the Cuban and 
Jamaican races of the species. 


Strabala ferruginea (Olivier) 
Ficure 17, 7 


Altica ferruginea Olivier, Entomologie .. . , vol. 6, p. 697, 1808. 
Strabala ferruginea Chevrolat, in d’Orbigny, Dictionnaire universel d’histoire 
naturelle, vol. 12, p. 52, 1848. 

Olivier’s description of the scutellum of this species as ferrugineous 
would differentiate it from the rest of the West Indian species even if 
he had not given the type locality as Santo Domingo. He described 
the color of the undersurface as black and this agrees with many of 
the specimens, but there is a certain variation in the coloring, some 
specimens being entirely ferrugineous beneath, others with dark legs 
and reddish breast and abdomen, others with dark legs, breast, and 
abdomen. Moreover the pronotum and elytra are finely and a little 
more distinctly punctate than in Cuban and Jamaican species and 
not at all alutaceous. The aedeagus, having a truncate tip, resembles 
the Puerto Rican specimens more than the Cuban or Jamaican. 


CHRYSOMELID GENUS STRABALA CHEVROLAT—BLAKE 133 


Distribution.—Haiti: La Vanneau, June 1920, Bizotan, Nov. 3, 
1925, Diguini, Apr. 14, 1925, all collected by W. A. Hoffman; Grande 
Riviére, W. M. Mann; Ile de la Tortue, April 1929, E. C. and G. M. 
Leonard; Port-au-Prince and vicinity, October 1934, P. J. Darlington 
(MCZ); Miragoane, October, November, P. J. Darlington (MCZ); 
Carrefour, May 1-3, 1908, M. Cameron (British Museum). Dominican 
Republic: August Busck; Macoris, Mar. 26, 1913, P. G. Russell; Puerto 
Plata, Hurst (MCZ); Santiago, 1938, P. J. Darlington (MCZ) 
Sanchez, July 1938, P. J. Darlington (MCZ). 


Strabala intermedia Jacquelin Du Val 
Figure 17, 1 


Strabala intermedia Jacquelin Du Val, in Ramén de la Sagra, Historia . . . de la 
Isla de Cuba (Spanish ed.), vol. 7, p. 129, 1857. 

In coloring, three old specimens in the Museum of Comparative 
Zoology labeled “Cuba’’ resemble Jacquelin Du Val’s short description. 
They are deep yellowish brown with dark antennae, dark scutellum, 
dark tibiae and tarsi; the anterior femora and the apex of the pos- 
terior femora are dark. In size they are slightly larger than the 
measurements given for S. intermedia; instead of being 3% mm. they 
are 4.5 to 5.5 mm. in length. In two, both females, the basal thoracic 
sulcus is pronounced, almost like that of a true Lactica, but in the third 
specimen, a male, the depression along the base is less marked. The 
surface is faintly alutaceous and finely punctate. In all three speci- 
mens the head is rather densely punctate in addition to the circle 
of depressed punctures near the eye. The coloring of these specimens 
is more like that of the North American species. The aedeagus, 
although essentially the same as in all the others, is distinctive in 
being slightly constricted behind the tip as in the Mexican and Central 
American species S. rotunda, but differs in having a more acute tip. 


Strabala colombiana, new species 


Figure 17, k 


From 5 to 6 mm. in length, oblong oval, deep reddish brown, the 
antennae, breast, abdomen, and legs dark, shining, not alutaceous, 
very finely, and the elytra rather densely, punctate. 

Head usually entirely deep reddish brown, shining, with scattered 
fine punctures over occiput and the usual depressed circle of punctures 
or fovea near eye. Antennae short, dark, the two basal joints paler. 
Prothorax shiny, finely punctate, a depressed line near base. Scutel- 
lum dark. Elytra shiny, very finely and indistinctly punctate. Body 
beneath with breast and abdomen deep piceous, legs dark. Length 
5 to 6 mm.; width 2.8 to 3.2 mm. 


134 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Type.—Holotype male, in Museum of Comparative Zoology, and 
1 paratype, USNM 61209, from Bogota, Colombia, from the Bow- 
ditch collection, Jacoby’s second collection. 

Other localities Colombia; Nueva Granada; Venezuela; and Cara- 
cas; all in the Bowditch collection, Jacoby’s second collection. 

Remarks.—Among the material presented to the U. S. National 
Museum from the Bowditch collection is a single old specimen 
of Strabala sp. with the label ‘‘Zactica=Strabala haematina Dej’’ 
from ‘Nov. Granad.’”’ This may be one of the specimens listed in 
the Dejean Catalogue from Cartagena since it bears the old Dejean 
Catalogue name. The aedeagus of this old specimen has a rounded 
tip quite unlike the one from Bogota with its acute tip, and is probably 
not the same species. 


Strabala trinitatis, new species 


Ficure 17, m 


From 4 to 4.9 mm. in length, oval, shiny, not alutaceous, finely 
but distinctly punctate, yellowish or reddish brown with darker 
antennae, scutellum, legs, breast and abdomen. 

Head entirely reddish, shiny, finely punctate over occiput. Anten- 
nae dark, with the first two or three joints paler. Prothorax shiny, 
finely punctate, with a moderately developed sulcus over the scutel- 
lum. Elytra distinctly and rather densely punctate, shiny, not at 
all alutaceous. Body beneath with breast, abdomen, and legs deep 
reddish brown. Length 4 to 4.9 mm.; width 2.5 to 2.7 mm. 

Type.—Holotype and 4 paratypes, British Museum, and 1 paratype, 
USNM 61210, collected in Trinidad in 1903 by G. E. Bryant. 

Other localities —Two specimens (USNM) collected by A. Busck on 
June 12 at Montserrat, Trinidad. 

Remarks.—The specimens in this series, although of the coloration of 
the ones from Colombia and Venezuela, are smaller, more oval, and 
distinctly though finely punctate. The tip of the aedeagus is truncate 
and not at all acute as in the single male specimen from Bogotéa. 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


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SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1953 No. 3320 


AMERICAN BITING MIDGES OF THE HELEID GENUS 
MONOHELEA 


By Wiis W. Wirt’ 


The genus Monohelea was founded by Kieffer (1917) for a new 
species, hieroglyphica, from Paraguay, and three other American 
species were included: Ceratopogon sequax Williston, 1896; C. maculi- 
pennis Coquillett, 1905; and C. nebulosa Coquillett, 1901. Since then 
a number of species have been added to the genus, including repre- 
sentatives from each of the world’s zoogeographic regions. However, 
with the exception of Lee’s (1948) treatment of the eight Australasian 
species, no comprehensive paper on the genus has yet appeared. 

A study of the American Monohelea has resulted in the recognition 
of 7 undescribed species, bringing the total for the hemisphere to 13. 
No doubt many more await discovery. These American species fall 
in four distinct groups, each with characteristic wing and leg markings. 
Within these groups the species are rather difficult to separate, for the 
characters which prove useful in one group may be valueless in others. 
These groups may also be recognized by characters of the male geni- 
talia, which offer in addition good specific characters. 

The generic position of Ceratopogon sequar Williston, based on a 
male from St. Vincent, cannot be determined with certainty until 
the male genitalia or the female are studied. According to the original 
description, seguax would probably fit as well in Stilobezzia. Simi- 
larly, the male recorded as Monohelea sp. by Floch and Abonnenc 
(1942) from French Guiana could be either Monohelea or Stilobezzia, 
since in their figure of the male genitalia the aedeagus is omitted. 

The types of five of the new species, as well as most of the specimens 
studied, are in the U.S. National Museum. For the loan or donation 

1 Bureau of Entomology and Plant?Quarantine, Agricultural Research Administration, U. S. Depart- 
ment of Agriculture. 
23299253 135 


136 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


of other specimens examined I am deeply grateful to Paul Freeman, 
of the British Museum (Natural History), to Henry Dietrich, of 
Cornell University, and to John Lane, of the University of S40 Paulo. 
For the generous gift or loan of still other specimens I wish to thank 
Mrs. Elisabeth C. Beck and J. A. Mulrennan of Jacksonville, Florida, 
Bernard Brookman of San Francisco, California, Jean A. Laffoon of 
Ames, Iowa, and Henry K. Townes of Raleigh, North Carolina. 

The figures of the wings and legs were made with the aid of a micro- 
projector and those of the male genitalia with the aid of an ocular 
reticule disc and squared paper. No consistent scale was used. 


Family HELEIDAE 
Genus Monohelea Kieffer, 1917 


Diagnosis —Body short, not very hairy. Eyes bare. Mesonotal 
pits present but small. Forelegs and midlegs unmodified; fourth 
tarsal segments cylindrical and claws small and equal in both sexes; 
hind leg with femur and tibia more or less thickened, but without 
spines, basitarsus with a spine at tip, fourth segment cylindrical, fifth 
segment in female with one very long claw, the other absent or very 
short, male claws small and equal except hind claw as in female in 
tessellata group; empodium absent. Wings with fine microtrichiae, a 
few macrotrichiae at wing tip; costa extending well beyond middle of 
wing; two radial cells, second longer than first; intercalary fork fairly 
distinct; crossvein r-m vertical; median fork with short stem, M2 
sometimes interrupted at base; anal vein thickened in middle, but 
without fold at thickening. Male genitalia with ninth tergite tapered, 
the caudal margin truncate or broadly bilobed; aedeagus usually with 
a pair of sharp-pointed, tapered, lateral sclerites and an accessory, 
dorsomedian, posterior lobe; parameres consisting of paired, sub- 
median sclerites of irregular shapes. 


Key to the American species of Monohelea 


1. Wings without markings; mesonotum shining black (subgenus Schizohelea). 
1. leucopeza (Meigen) 
Wings with pronounced dark markings; mesonotum pruinose, often with mark- 


ings (subgenus Monohelea)_..--.5----. 23.2222. 2 ee Ls 2 

2. Hind femur and tibia yellow, with dark brown rings (hieroglyphica group)- 3 
Hind femur and tibia shining black, only knee spot yellow__-_----------- 9 

3. Forelegs and midlegs with distinct dark brown bands; wings often with exten- 
sive mserotrichiae..__.41.. ~~ -- 52. 4 
Forelegs and midlegs yellowish, without distinct bands; wings with macro- 
trichiae only at extreme distal margin. -_--_.__.....-==22 So eee 6 

4. Wings with macrotrichiae over distal third; wing markings extensive__._ 5 


Wing without apparent macrotrichiae; wing markings reduced, X-shaped 
marking in cells M; and Mz broken up into 3 discrete spots. 
6. texana, new species 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 137 


5. Wing with about 20 small, scattered, black dots in addition to the irregular, 
diffuse gray markings; fore tibia and midtibia dark only at apices; scutellum 
TMIfO ceva wilifas ye yee age ee es ke 8. brasiliensis Lane 

Wing with uniformly gray, irregular markings, without small, black dots; 
fore tibia and midtibia entirely dark; scutellum dark in middle. 
7. ornata, new species 

6. Hind femur brown on basal third; scutellum dark in middle; wing markings 
Asta HI bis bOndere deme a = eeyr ee sie ee So ape ee eee 7 

Hind femur with broad yellow band on sub-basal fourth; scutellum uniformly 
dull whitish; wing markings bordered with whitish pruinosity. 
3. maculipennis (Coquillett) 

7. Basal and median brown bands on hind tibia joined with brown, at least on 
ventral side; male parameres each with 2 or 3 distal lobes______------- 8 

Basal and median brown tibial bands separated by a broad yellow band; male 
parameres each with simple, bladelike apex__-_-- 5. macfiei, new species 

8. Sinuate, dark, wing marking in cells M; and Mg entire; male parameres with 
prosds rounded! distal) lobesss22252-)- 934555522 4. lanei, new species 

Sinuate, dark, wing marking broken into 3 or 4 separate spots; male parameres 
with slender, pointed, distal lobes__________-_- 2. hieroglyphica Kieffer 

9. Scutellum yellow, brown in middle; wing with 2 prominent, small, black spots, 
spot absent in basal cell; hind basitarsus about 2.5 times as long as second 
segment; mesonotum yellowish, with tiny brown dots (multilineata group) 10 

Seutellum entirely dark brown or yellow only in middle; wing with two prom- 
inent anterior spots plus a spot in basal cell; hind basitarsus 1.3 to 2.0 times 


as long as second segment (tessellata group)____-_----------------- 12 
10. Ninth sternite of male with 4 long hairs arising from tubercles in a row at base 
GMIMeECIANV OMe was! a a! See ee Bee ee. eet hee 11 


Ninth sternite of male with 2 long hairs arising from base of median lobe; para- 
meres not connected, with irregular, twisted apices; female abdomen 
pruinose gray above, with small, lateral, shining brown spots. 

9. stonei, new species 

11. Male aedeagus with broad, bilobed apex; parameres not connected, recurved 

apices long and scimitar shaped; female abdomen pruinose gray above, with 
narrow, shining brown bands broadening at margins of segments. 

10. multilineata (Lutz) 

Male aedeagus cleft one-third way to base, with a pair of long, slender, apical 
points; parameres broadly connected at bases, recurved apices with broadly 
ROUNGEGOeOWESE {2 a Bake bel es 11. guianae, new species 

12. Mesonotum pruinose gray with dark-brown spots or irregular patches; scutel- 
lum yellow in middle, with about 8 marginal hairs; wing bands distinct to 
hind margin, usually strongly interconnected___ 12. nebulosa (Coquillett) 

Mesonotum dark brown with irregular, pruinose, gray spots; scutellum usually 
entirely black, with 4 marginal hairs; wing bands faint behind vein My, 
usually rather well separated__....._.._---- 13. johannseni, new species 


Subgenus Schizohelea Kieffer, 1917 


Schizohelea Kieffer, Ann. Mus Nat. Hungarici, vol. 15, p. 295, 1917, vol. 16, p. 57, 
1918, and vol. 17, p. 89, 1919.—Goetghebuer, Mem. Mus. Hist. Nat. 
Belgique, vol. 8, p. 65, 1920.—Edwards, Trans. Ent. Soc. London, vol. 74, 
p. 411, 1926—Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 782, 1943. 
(Genotype, Ceratopogon copiosus Winnertz; monobasic.) 

Allohelea Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 364, 1917. (Genotype, 
Sphaeromyas pulchripennis Kieffer; origina] designation.) 


138 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Following Goetghebuer (1934), I regard Schizohelea as a subgenus 
for leucopeza Meigen. 

Diagnosis.—This species differs rather markedly from the other 
species of Monohelea in having the second branch of the media broadly 
interrupted at the base, basitarsus without apical spine, and male 
genitalia with emarginate sternite and arched aedeagus. 


1. Monohelea (Schizohelea) leucopeza (Meigen), 1804 
FicurE 18, m 


Ceratopogon leucopeza Meigen, Klassifikation und Beschreibung . . . Insekten, 
vol. 1, p. 29, 1804. 

Ceratolophus leucopeza, Kieffer, Genera insectorum, fasc. 42, p. 60, 1906. 

Schizohelea leucopeza, Edwards, Trans. Ent. Soc. London, vol. 74, p. 411, 1926.— 
Séguy, Faune de France, pt. 8, p. 70, 1937. 

Monohelea (Schizohelea) leucopeza, Goetghebuer, in Lindner, Die Fliegen der 
Palaearktischen Region, Lief. 78, p. 53, 1934. 

Ceratopogon albitarsis Wiedemann, Zool. Mag., vol. 1, p. 67, 1817. 

Ceratopogon copiosus Winnertz, Linn. Ert., vol. 6, p. 56, 1852. 

Ceratolophus copiosus, Kieffer, Genera insectorum, fasc. 42, p. 60, 1906. 

Schizohelea copiosa, Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 295, 1917; idem, 
vol. 16, p. 57, 1918; idem, vol. 17, p. 89, 1919.—Goetghebuer, Mem. Mus. 
Hist. Nat. Belgique, vol. 8, p. 66, 1920.—Kieffer, Faune de France, pt. 11, 
p. 116, 1925. 

Ceratopogon politus Coquillett, Proc. U. S. Nat. Mus., vol. 23, p. 606, 1901 (9, 
Massachusetts). 

Ceratolophus politus, Kieffer, Genera insectorum, fasc. 42, p. 61, 1906. 

Johannseniella polita, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 
227, 1914. 

Johannsenomyia polita, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 
335, 1915 (New York). 

Sphaeromias polita, Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 364, 1917. 

Allohelea polita, Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 364, 1917. 

Schizohelea polita, Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 782, 1943 
(? =leucopeza Meigen). 

Bezzia stecki Kieffer, Brotéria, Ser. Zool., vol. 18, p. 65, 1915. 

Description —FeEmate: Length 1.5 mm., wing 1.2 mm. by 0.5 mm. 
Entirely shining black, only the tarsi, wings, and halteres white. 
Antennae quite short, the distal segments scarcely elongated. 
Mesonotum with scattered, erect, long, black hairs; scutellum with 
six long marginal hairs. Second radial cell half again as long as the 
first, Mz broadly interrupted at base. Spermathecae two, subequal, 
rather small and subspherical, the ducts sclerotized a short distance. 

Matz (based on specimen from Suffolk, England): Ninth sternite 
about three times as broad as long, with a shallow, round emargination 
in middle of caudal margin, not spiculate; ninth tergite narrow, sur- 
passing basistyles, distal half with margins subparallel, apex truncate 
with setose apicolateral corners, the inner surface coarsely spiculate. 
Basistyles broad at base, each with a distinct lobe bearing three setose 
tubercles on inner side; abruptly narrowed on distal half; dististyles 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 139 


TABLE 1.—Proportions of segments of hind legs of female Monohelea species 


Species Cx AME F | aay Ay Te T3 Aly; Ts 
i leucopera.o- |=). 2- 25 10 65 65 30 12 10 8 12 
2. hieroglyphica_-_-___- 30 10 80 | 72] 40 16 9 8 8 
3. maculipennis_ -__-_-__- 20 10 55 50 24 11 8 6 + 
cS 0) Ses ee ae 25 10 65 60 | 35 17 10 8 8 
Se 20 10; 60); 55 30 15 10 7 7 
perorentae 24 St? 72) 681" 3a 16 9 rs 7 
Piepanet rt js esse 30 10 80 | 70] 50 20 12 8 10 
10. multilineata_______-_ 28 12 | 88 80 50 20 12 8 8 
fe penuloss-—_...___.- 28 12 | 80; 70 24 12 6 8 8 
i32-johannseni. 222+. 25 10 70 65 25 12 8 10 10 


nearly as long as basistyles, rather strongly curved. Aedeagus in 
form of a slender, rounded arch nearly as long as broad at base, with 
a pair of short, pointed, submedian plates projecting dorsocaudad 
at apex. Parameres with stout, lateral apodemes at bases; stems 
stout and gently curved, contiguous at midlength, their apices pointed 
and abruptly bent ventrocephalad on about distal fourth. 

Type.—Presumably in the Meigen collection in the Muséum 
National d’Histoire Naturelle in Paris. 

Material examined.—Massachusetts: N. Amherst, June 1940, 4 99. 

New York: Mecklenburg, June 2, 1940, A. Stone, 10 99; North 
Ridgeway, June 22, 1940,S. C. Mendall, 1 9. 

Michigan: St. Joseph, May 30, 1938, C. W. Sabrosky, 6 99. 

England: Corriegills, Arran, June 2-4, 1919, F. W. Edwards, 2 99; 
Mildenhall, Suffolk, May 22, 1909, Yerbury, 1 . 

Remarks.—This species is found in Europe, West Africa, and 
northeastern North America. 


Subgenus Monohelea Kieffer, 1917 


Monohelea Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 295, 1917.—-Goetghebuer, 
Mem. Mus. Hist. Nat. Belgique, vol. 8, p. 63, 1920.—Ingram and Macfie, 
Ann. Trop. Med. Parasit., vol. 15, p. 344, 1921.—Edwards, Trans. Ent. Soc. 
London, vol. 74, p. 410, 1926.—Tokunaga, Tenthredo, vol. 3, p. 156, 1940.— 
Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1948—-Lane, Arq. Fac. Hig. 
Saude Univ. Saéo Paulo, vol. 1, p. 225, 1948.—Lee, Proc. Linn. Soc. New 
South Wales, vol. 72, p. 350, 1948. (Genotype, Monohelea hieroglyphica 
Kieffer; original designation.) 

Diagnosis.—Wings with prominent markings; body with pruinose 
pattern; ninth sternite of male spiculate and transverse or lobed, not 
emarginate. 

hieroglyphica—group 


Diagnosis.—Hind legs conspicuously banded, not markedly swollen; 
hind tarsi slender; wings with extensive irregular markings; male 
aedeagus broad, with median, anterior notch and triangular, lateral 
sclerites with converging, sharp-pointed apices. 


140 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


2. Monohelea (Monohelea) hieroglyphica Kieffer, 1917 


FicureE 19, g 
Monohelea hieroglyphica Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 312, 1917 
(a, 9, Paraguay). 
Monohelea hyeroglyphica, Lane, Arq. Fac. Hig. Saude Pub. Univ. Sao Paulo, vol. 
1, p. 225, 1948 (Brazil). 

Discussion —This is the genotype of Monohelea. Macfie’s (1937, 
1940a) records of this species from Trinidad and British Guiana must, 
I believe, be referred to macfiet, new species, described (see. p. 143), 
from Louisiana. 

Through the kindness of John Lane I have examined a female from 
km. 47, estrada Rio-Séo Paulo, Rio de Janeiro, Brazil, collected in 
February 1945 by P. Wygodzinsky. This is from the same locality 
as the male described and figured by Lane. Externally this species is 
almost inseparable from M. (M.) lanei, new species (see p. 142), from 
Florida, but the male genitalia are very close to those of M. (M.) 
maculipennis. A sketch of the male parameres (after Lane) is included 
here for comparison with those of maculipennis. 

Types.—In the Musei Nationalis Hungarici in Budapest, o’, 9, 
Paraguay. 


3. Monohelea (Monchelea) maculipennis (Coquillett), 1905 
Fiaures 18, a, 7; 19, f 


Ceratopogon maculipennis Coquillett, Journ. New York Ent. Soe., vol. 13, p. 64, 
1905 (9, Florida).—Kieffer, Genera insectorum, fase. 42, p. 51, 1906. 

Monohelea maculipennis, Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 312, 1917.— 
Johannsen, Ann. Ent. Soc. Amer. vol. 36, p. 781, 1948. 

Diagnostic characters—A yellowish gray, pruinose species with 
faint, brown, mesonotal dots, pale yellowish scutellum; hind legs with 
narrow, dark rings and wings with extensive, irregular, grayish macula- 
tions bordered with whitish pruinosity. 

Description.—Frmate: Length 1 mm., wing 1 mm. by 0.4 mm. 
Head pale yellowish, antennae and palpi brown; proportions of flagellar 
segments of antennae 12:10:10:12:12:12:12:12:16:16:18:18:24. 
Palpal segments in proportion of 5:7:10:5:10. 

Mesonotum grayish pruinose, with many scattered, small, brown 
dots at the bases of the dark mesonotal hairs; humeri and sides exten- 
sively yellow. Scutellum pruinose, yellowish white, with four marginal 
hairs, the middle pair quite close together. Postscutellum and pleura 
pruinose brown, the latter with several indistinct, transverse, darker 
lines. Legs yellowish, coxae and trochanters brown, upper fourth of 
midcoxa and hind coxa yellowish; fore femur and midfemur and tibiae 
unbanded; hind femur (figure 18, 7) brown at extreme base, two narrow 
oblique dark rings in middle and a dark, preapical, ventral spot; hind 
tibia with narrow sub-basal, median, and apical rings; tarsi narrowly 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 141 


dark at apices of segments. Proportions of segments of hind legs as in 
table 1. 

Wing whitish hyaline, with irregular maculations, grayish by 
transmitted light, yellowish brown with narrow, whitish pruinose 
borders by reflected light. Markings composed of a broad, irregular 
band across wing at level of first radial cell, filling basal half of medio- 
cubital fork, with a prominent, omega-shaped spur in base of cell R;; 
a small spot before middle of second radial cell; a more or less quadrate 
to X-shaped mark across cell R; at apex of second radial cell; a sigmoid, 
subapical mark across cells M, and Mz, often connected by very narrow 
lines in these cells to mesal band; small, irregular spots across wing near 
base and a small spot past middle of basal cell. First radial cell about 
half as long as second, a very few macrotrichiae at apices of cells R; 
and M,. Halteres dull white, a black dot on anterior side. 

Abdomen dull whitish, sides with irregular, dark patches. Sper- 
mathecae two, very unequal, subspherical, each with short sclerotized 
duct. 

Maus: Ninth sternite about twice as broad as long, spiculate, 
posterior margin transverse, abutting against base of aedeagus, with 
four long hairs arising from tubercles in a curved row; ninth 
tergite greatly constricted, with sides subparallel on distal half, apex 
truncate, apicolateral processes short. Basistyles narrowed on distal 
halves; dististyles curved to slender, pointed apices. Anterior margin of 
aedeagus broad, slightly concave, with a small median notch; aedeagus 
bearing a pair of triangular, submedian sclerites, the bladelike apices 
of which converge before level of the irregular bilobed apices of an 
accessory pair of dorsal sclerites. Parameres with flaring, winglike, 
bilobed, basal apodemes, more or less connected at midlength by a 
pair of stout mesal lobes, apices each with a straight, long, slender 
stem bearing a dorsolateral lobe about half as long and of same thick- 
ness, just beyond level of mesal bridge. 

Type—USNM 8366, 9, Florida, Jacksonville. 

Material examined.—Florida: Crystal River, Citrus County, July 
17, 1950, Hudson, 4 99; Everglades City, Collier County, Feb. 7, 
1950, Davidson, 1 o, 9 99; Fort Myers, Lee County, Feb. 2, 1949, 
Brechtel, 1 &; Islamorada, Monroe County, June 7, 1949, Smith, 1 
o, 2 99; Jacksonville, date not given, Slosson, 1 9 (type). 

México: Ciudad Monte, Tamaulipas, Nov. 22, 1943, Brookman, 1 
BY S-99. 

Guatemala: Rio Dulce, Mar. 21, 1906, Schwarz and Barber, 1 9. 

Panamé: Rio Trinidad, June 9, 1902, Busck, 1 9. 

Remarks.—¥ormerly all the North American Monohelea with the 
type of wing pattern characteristic of hieroglyphica were called 
maculipennis. Thus in my (1952) paper on California Heleidae, I 


142 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


erroneously included a California record of maculipennis and also 
cited several eastern records of this speciesin error. I have now 
studied the hieroglyphica group more intensively and believe I have 
correctly identified several component species on the basis of male 
genitalia supported by several rather difficult but constant external 
characters of the female. In addition to hieroglyphica and maculi- 
pennis there appear to be four or more undescribed species in this 
complex. Although my California specimen and that reported as 
Monohelea sp. by Johannsen (1943) do not fit any of those described 
below, probably falling closest to hieroglyphica in wing markings, a 
positive diagnosis of the species must await the collection of the male. 


4. Monohelea (Monohelea) lanei, new species 
Fiaures 18, b, j; 19, e 


Description.—FEMALE: Length 1.3 mm., wing 1.1 mm. by 0.4 mm. 
As in M. (M.) maculipennis (Coquillett), but darker and slightly 
larger. Mesonotum light brown pruinose with faint brown punctures; 
scutellum brownish in middle. Markings on hind legs stronger; 
basal third of hind femur brown, a narrow, diagonal band just past 
middle, then a small, brown, ventral spot and a strong preapical band; 
basal and median bands on hind tibia broad and joined on ventral side, 
the distal band broad. Proportions of segments of hind leg as in 
table 1. 

Wing markings darker and more extensive than in maculipennis, 
the omega-shaped mark entirely closed behind by a broad extension 
of the mesal band, the X-shaped mark in cell R; greatly constricted in 
middle and much broader behind. Macrotrichiae sparse and con- 
fined to wing margin; first radial cell about two-thirds as long as 
second. 

Abdomen brown above, yellowish on sides in front and on three 
distal segments. Spermathecae two, large, very unequal and sub- 
spherical. 

Maus: Ninth sternite with posterior margin convex in middle, 
spiculate; ninth tergite tapered to a pair of blunt, triangular, sub- 
median, caudal lobes. Basistyles narrow, nearly straight; dististyles 
slender, curved, with sharp, incurved points. Aedeagus with basal 
halves broadly separated by a deep notch in middle of anterior margin, 
lateral sclerites with broad, triangular bases, the slender, pointed, 
posterior portions directed caudomesad and meeting each other over 
tips of a pair of sharp-pointed, dorsal processes. Parameres a pair 
of irregular, sublateral sclerites joined by an indistinct transverse 
bridge at a third of the distance from bases; basal apodemes trilobed, 
directed laterocephalad to bases of basistyles; posterior portions with 
subparallel inner margins, tips each with apical, knoblike, sclerotized 
lobe with slightly longer, flattened, roundly flaring, ventral expansions. 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 143 


Types.—Holotype, o&, USNM 61091, Miami, Florida, Mar. 25, 
1944, Wirth. Allotype with same data as type except date, October 
1943. Paratypes: Florida: 2 7,599, same, except date, October 
1943 to April 1944; 1 9, Welaka, June 17, 1946, Bellamy; 1 9, 
Everglades City, Feb. 7, 1950, Davidson. 


5. Monohelea (Monohelea) macfiei, new species 


Figures 18, c, h; 19, h 
Monohelea hieroglyphica, Macfie, not Kieffer (misidentification), Ann. Mag. Nat. 
Hist., ser. 10, vol. 20, p. 18, 1937 (?, Trinidad); Ent. Monthly Mag., vol. 
76; p. 80, 1940 (0, British Guiana, genitalia figured). 

Description.—FEMALE: Length 1.2 mm., wing 1.4 mm. by 0.5 mm. 
As in M. (M.) maculipennis (Coquillett), but the mesonotum with a 
pronounced, broad, median, anterior band and a large area between 
wing bases in front of scutellum uniform brown, the scutellum with a 
more or less brownish median area. Abdomen suffused, grayish 
brown above. Legs as figured by Macfie (1937), brown bands on 
hind legs quite distinct; on hind femur a band on basal third, a diagonal 
band past middle, and a narrow band subapically;on hind tibia, 
narrow basal, median, and apical rings; all tarsal segments with 
narrow apical bands. 

Wing as in figure 18, c, the dark areas in the first basal and anal cells 
quite extensive, the X-shaped mark in cell R; large, with the caudo- 
distal arm more prolonged than shown by Macfie and the distal sig- 
moid mark in cells M, and M, connected by broad lines in these cells 
to the broad median crossband. All wing markings brown by re- 
flected light, gray by transmitted light; a few macrotrichiae at apex 
of wing. 

Mate: The resemblance of the Louisiana male to the male from 
British Guiana figured by Macfie (1940a) is remarkable. The 
slender, sinuate, bladelike parameres connected by a narrow bridge 
at midlength are the most distinctive difference. 

Types.—Holotype, o', USNM 61092, allotype, Kilbourne, Louisiana, 
May 10, 1947, Wirth. Paratypes: Louisiana: 2 9? , with same data 
as type; 1 9, Baton Rouge, May 16, 1947, Wirth. 


6. Monohelea (Monchelea) texana, new species 


Ficure 19, d 


Description —Matz: Wing 1.1 mm. long. Badly damaged, anten- 
nae, all of legs but fore feraur, midfemur, and tibiae gone. Mesonotum 
appears discolored, all dark except large quadrate yellow patches on 
humeri; scutellum light yellow, a minute brown spot in middle of 
anterior surface. Forelegs and midlegs yellow, narrow basal, median, 
and subapical faint brown rings on fore femur; narrow basal and 

232992532 


144 PROCEEDINGS OF THE NATIONAL MUSEUM bh. 


broader median and apical brown rings on fore tibia; broad basal and 
narrow median brown rings on midfemur and faint median and 
broad, dark-brown apical rings on midtibia. Wings marked as in 
M. (M.) hieroglyphica Kieffer, the median band broad, the omega- 
shaped spur distinct and open below, the spot in cell Rs narrow with 
the right-hand arms of the X absent, the sinuate mark in cells 
M, and M-z reduced to three discrete spots; macrotrichiae reduced to 
a few at wing margin. Halteres yellow with flat ends and a spot on 
anterior surfaces of knobs dark. 

Ninth sternite spiculate, posterior margin broadly convex in middle, 
with four long hairs arising from bases in an arched row; ninth tergite 
tapered, with a pair of prominent, apicolateral processes. Aedeagus 
with basal halves narrowly separated by a deep notch in middle of 
anterior margin, lateral sclerites with broad, triangular bases, the 
slender, pointed posterior apices meeting each other at about level 
of tip of ninth tergite; the dorsal accessory structure bifid a third 
the way to base of aedeagus, with a pair of very slender lobes with 
pointed apices curved ventrad and surpassing apices of lateral sclerites. 
Parameres joined together broadly near middle, each with well- 
developed basal and lateral arms, stems abruptly bent laterad two- 
thirds the way to apices, then abruptly bent ventrad, with pointed 
apices directed mesocaudad. 

Type.—Holotype, o', Cornell Univ. Type 2787, Limpia Canyon, 
Davis Mountains, Texas, July 7, 1917, Bradley. 

Remarks —The male genitalia are most distinctive, the simple 
apices of the parameres allying M. (M.) terana with M. (M.) macfiei, 
new species. However, the external features, as nearly as can be 
ascertained from the badly damaged specimen, are closer to other 
species, the wing markings being nearly like those of M. (M.) hero- 
glyphica Kieffer, while the front legs and midlegs are banded as in 
M. (M.) ornata, new species, and M. (M.) brasiliensis Lane. 


7. Monohelea (Monohelea) ornata, new species 
Fiaure 18, d 


Description.—FEMALE: Length 1.2 mm., wing 1.2 mm. by 0.5 mm. 

Head pruinose brown, vertex yellowish; antennae broken, pedicel 
brown; palpi brown. Mesonotum pruinose gray, humeri yellowish; 
a broad, median, longitudinal band and broad patches above wings 
rich, velvety brown. Scutellum pruinose, grayish yellow, brown in 
middle; postscutellum and pleura pruinose dark brown. Fore femur 
and midfemur dull yellow, with faint median brown bands; fore tibia 
and midtibia brown; hind femur dark brown on basal third, a broad, 
diagonal, brown ring past middle and a narrow preapical brown ring; 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 145 


hind tibia with broad basal and apical rings, and a band just before 
middle, brown; tarsi dull yellowish. Proportions of segments of 
hind legs as in table 1; hind femur and tibia with rather long hairs; 
hind basitarsus with strong basal spine. Claws on forelegs long and 
equal, nearly as long as fourth and fifth segments combined; midtarsi 
broken ;on hind leg a single long claw half again as long as fifth segment. 

Wing grayish hyaline, with very extensive gray maculations as in 
figure; maculations of hieroglyphica type, but much more extensive, 
with an extra distal, separate spot narrowly connected to the X-shaped 
mark in cell Rs, and the subapical mark in cells M; and Mz quite broad. 
First radial cell about half as long as second; macrotrichiae very ex- 
tensive, sparsely covering distal third of wing and including most of 
cell M ;4 4 and anal cell. Halteres not visible. Abdomen uniformly 
dark, pruinose brown, spermathecae not examined. 

Types.—Holotype, 2, USNM 61093, Santa Rosa Island, Escambia 
County, Florida, Oct. 10, 1949, Butler. 

Remarks.—The wing, mesonotal, and leg markings are so distinctive 
that I do not hesitate to describe M. (M.) ornata from the single female. 


8. Monohelea (Monohelea) brasiliensis Lane, 1948 


Monohelea brasiliensis Lane, Arq. Fac. Hig. Saude Pub. Univ. Sao Paulo, vol. 1, 
p. 226, 1948 (9, Brazil). 

Description.—I have not seen this species, which is known only from 
the type female. Length 1.2 mm., wing 1.3 mm.; mesonotum chest- 
nut, yellowish on sides and in prescutellar depression; scutellum 
whitish. Legs yellowish, fore femur dark at base, midfemur dark on 
basal half; fore tibia and midtibia dark at apices; hind femur with two 
dark rings, one at apex of basal third and one in middle, hind tibia dark 
at base and apex and indistinctly so in middle. Hind basitarsus 1.8 
times as long as second segment. Wing with about 20 black dots in 
addition to irregular grayish markings; macrotrichiae numerous on 
distal third of wing; first radial cell slightly over half as long as 
second (from original description). 

Type.—In collection of University of Sao Paulo, Brazil, No. 6781, 
9, Brazil, Rio de Janeiro, Estrada Rio-Sdo Paulo, km. 47. 


multilineata—group 


Diagnosis.—Wings with two large anterior dark patches and other 
fainter irregular markings; scutellum dark in middle; Jegs dark with 
yellow knee spots; hind legs slender, the tarsi long and unspined; 
ninth sternite of male with median, convex lobe and several long 
hairs, aedeagus long and narrow, parameres very long, with sharp 
apices. 


146 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


a 
D 


b. LANE] 


, Any 


g JOHANNSENI 


d.ORNATA 


VQ 


i 
JOHANNSENI | 


h. MACFIEI 


j- LANE! k. JOHANNSENI. m.LEUCOPEZA 


Ficure 18.—Species of Monohelea: a-g, Wings: a, maculipennis (Coquillett); b, Janet, new 
species; ¢, macfiet, new species; d, ornata, new species; ¢, stone1, new species; f, nebulosa 
(Coquillett); g, johannseni, new species. h-k, Hind legs of females: h, macfies, new species; 
4, maculipennis (Coquillett); 7, lanei, new species; k, johannseni, new species. 1, Sperma- 
thecae, johannseni, new species. m, Male genitalia, leucopeza (Meigen). 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 147 


Mi! $y 
E ie 


h 


YA 


i‘ 


fpae 
WA 


a. JOHANNSENI 


¢. LANEI 


d. TEXANA 
g- HIEROGLYPHICA 


i. STONE! j-MULTILINE ATA k, GUIANAE 


Ficure 19.—Species of Monohelea, male genitalia: a, johannseni, new species; b, tessellata 
(Zetterstedt); c, nebulosa (Coquillett); d, texana, new species; ¢, lanei, new species; f, 
maculipennis (Coquillett); g, hieroglyphica Kieffer; male parameres (after Lane); h, macfiei, 
new species; 1, stonet, new species; 7, multilineata (Lutz); k, guianae, new species. 


148 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


9. Monohelea (Monohelea) stonei, new species 
Fiaures 18, e; 19, 7 


Diagnostic characters—A yellowish gray, pruimose species with 
maculate wings; scutellum yellow with brown center; legs dark, 
abdomen dull gray pruinose above. 

Description.—FEMALE: Length 2.0 mm., wing 1.5 mm. by 0.7 mm. 
Head yellowish, vertex pruinose, pale gray; antenna with pedicel and 
bases of flagellar segments yellow, apices of short segments and all of 
long distal segments and palpi brown; proportions of flagellar seg- 
ments 15:10:10:10:10:10:12:12:18:18:18:18:24. Palpal segments 
in proportion of 5:8:15:10:15, third segment scarcely swollen, with a 
rather large sensory pit at midlength. 

Mesonotum yellowish gray pruinose, with tiny brown dots at the 
bases of the brownish mesonotal hairs. Scutellum yellow, brown in 
middle, with four moderate marginal hairs and a few fine setae. 
Postscutellum and pleura pruinose dark brown. Coxae brown; 
trochanters, femora, and tibiae light brown on forelegs and midlegs, 
black on hind legs; all knees prominently yellowish, tarsi yellow. 
Hind femur and hind tibia moderately thickened; basitarsus with a 
slender spine at base, distal tarsal segments on hind legs each with 
a fine apical spine; a single long claw on each hind leg. Proportions 
of segments of hind leg as in table 1. 

Wing grayish hyaline, with two prominent, black spots and fainter 
posterior infumation as figured; first radial cell about half as long as 
second. Halteres infuscated. 

Abdomen dark brown, entire dorsum pruinose gray except for a 
pair of small lateral spots and a pair of submedian sensory dots on 
each segment, polished brown. Spermathecae two, subequal, ovoid, 
each with a very short sclerotized duct. 

Maus: Similar to the female, with the usual sexual differences; 
antennal plume golden at base, dark brown toward apex. Ninth 
sternite three times as broad as long, with a narrow, median, poste- 
rior lobe bearing two long hairs extending into the concave base of the 
aedeagus; sternite spiculate on posterior half; ninth tergite as long as 
basistyles, tapered to the truncated apex, bearing a pair of short, 
rounded, apicolateral processes. Basistyles about twice as long as 
broad, simple, very convex on outer sides; dististyles about 0.7 as long 
as basistyles, nearly straight and stout to apices. Aedeagus with 
base as broad as the lobe of ninth sternite, a slight sub-basal swelling, 
a second swelling at midlength, with distal half cleft into a pair of 
attenuated, submedian, pointed blades attaining 0.8 length of basi- 
styles. Parameres consisting of a pair of very irregular processes, 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 149 


bases of each with a large anterior and lateral apodeme, stem rather 
stout and crooked, distal portion abruptly bent ventrocephalad, then 
greatly attenuated and bent ventrocaudad to a sharp point. 

Types.—Holotype, 2, USNM 61094, Baton Rouge, Louisiana, May 
16, 1947, Wirth. Allotype with same data as type except date, 
May 3, 1947. Paratypes: Louisiana: 60707, 3 99, with same data as 
type except date, May 6-20, 1947. Mississippi: 4 oo’, 10 99, Horn 
Island, June and July 1944, E. A. Richmond. Florida: 11 99, Gray- 
ton Beach, Walton County, May, Sept., 1949, Butler; 1 9, Santa 
Rosa, Walton County, July 19, 1949, Peterson; 8 oo, 21 99, In- 
nerarity Point, Escambia County, May, 1950, Rathert; 8 99, Santa 
Rosa Island, Escambia County, July 7, 1949, Butler; 5 99, Panama 
City Beach, Bay County, May 6, 1949, McElvey; 5 99, Crystal River, 
Citrus County, July 17, 1950, Hudson; 3 99, Fort Myers, Lee County, 
Oct. 8, 1948, Brechtel; 3 99, Everglades City, May, Oct., 1948, 1950, 
Huntoon; 1 o, 1 9, Big Pine Key, Monroe County, Apr. 1, 1950, 
Sermon. Georgia: 2 oo’, 4 99, Thomasville, May 15-30, 1949, 
Palmer. Tennessee: 1 9, Nashville, June 18, 1937, Adams. Iowa: 
1 9, Sioux City, July 15, 1950, Laffoon. Virginia: 3 oo, 6 99, Falls 
Church, July 8, 29, 1950, Wirth. Maryland: 4 99, Leeds, Dorchester 
County, July 10, 1907, Barber. New Jersey: 1 9, Newport, July 1, 
1937 (light trap). Costa Rica: 1 o&, Higuito, San Mateo, no date, 
Schild. Panama: 1 9, Fort Kobbe, Canal Zone, Aug. 21, 1950, 
Carpenter. 


10. Monohelea (Monohelea) multilineata (Lutz), 1914 
Figure 19, j 


Palpomyia multilineata Lutz, Mem. Inst. Oswaldo Cruz, vol. 6, p. 93, 1914 (Brazil). 
Monohelea multilineata, Johannsen, Ann. Ent. Soc. Amert., vol. 36, p. 781, 1943.— 
Lane, Rev. Ent., vol. 16, p. 368, 1945. 

Lane (1945) has very adequately redescribed this Brazilian species. 
It resembles M. (M.) stonez, new species, in general coloration and 
structure, the mesonotum yellowish with brown dots, the scutellum 
yellow with median brown spot and four marginal bristles, forelegs 
and midlegs yellowish, the hind femur and tibia black and scarcely 
swollen and hind tarsi unspined. The wings, however, in addition 
to the two anterior black spots, have smaller, rather strong spots at 
the wing margin in cells R; and M,, and across the apex of cell Mz. 
The polished, brown, lateral spots on the abdomen, which are small 
in stonei, are larger and narrowly connected across the anterior margin 
of each tergite in multilineata. 

The male genitalia of multilineata are of the same type as those of 
stonei, but there are four rather than two bristles at the base of the 


150 PROCEEDINGS OF THE NATIONAL MUSEUM Yon, 72 


lobe of the ninth sternite, the aedeagus is broader at the apex and 
only slightly notched rather than deeply cleft, and the parameres are 
slender and nearly straight with the apices abruptly bent and scim- 
itar-shaped. 

Material examined.—Brazil: Km. 47, Estrada Rio-Séo Paulo, Rio de 
Janeiro, Brazil, February 1945, Wygodzinsky, 2 oc, 2 99. 

Remarks.—Monohelea nigeriae Ingram and Macfie, 1922, known 
from the female from West Africa, is closely related to MZ. (M.) stonei 
and to M. (M.) multilineata, but according to the original description, 
the tarsi bear ventral spines, as in the tessellata group, and the sperma- 
thecae are unequal in size. 


11. Monohelea (Monohelea) guianae, new species 
Ficure 19, k 


Monohelea multilineata, Macfie, not Lutz (misidentification), Proc. Ent. Soc. 
London, Ser. B, vol. 9, p. 187, 1940 (o’, British Guiana). 


Macfie (1940) has given as satisfactory a description of this insect 
as was possible from the single male specimen available. This speci- 
men was borrowed from the British Museum for study and externally 
cannot be separated from either M. (M.) multilineata (Lutz) or 
M. (M.) stonei, new species. However the male genitalia are quite 
distinct and may be characterized as follows: Ninth sternite very 
short, with a low median lobe on caudal margin fitting against base of 
aedeagus, spiculose, with four long hairs arising from tubercles in a 
line at base of median lobe; ninth sternite very narrow, tapered to 
tip, with a pair of short, nipplelike apicelateral processes. Basistyles 
about twice as long as broad; dististyles nearly straight, with blunt, 
scarcely narrowed apices. Aedeagus very narrow, about 2.5 times as 
long as broad at base, the anterior arch very low, apex cleft about a 
third way to base, with a pair of very slender, submedian processes, 
from the bases of which a pair of slender, lateral arms curve around 
stems of parameres. Parameres broadly joined together at bases, 
each with a short anterior and lateral arm, stems slender, apex of each 
thickened and abruptly bent Jaterocephalad in a broadly rounded 
lobe about half as long as dististyle. 

Types.—Holotype, o, British Museum (Natural History) collec- 
tion, Mazaruni, British Guiana, Aug. 21, 1937, second growth (low 
forest), Richards and Smart. 


tessellata—group 


Wings with three large dark anterior spots and irregular posterior 
infuscated areas; scutellum often with middle yellow; hind legs swollen 
and black, with yellow knees, the tarsi short and with strong ventral 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 151 


spines; claw of hind leg of male long and single as in the female; male 
aedeagus with slender, pointed, posterior sclerite borne in a notch in 
the quadrate basal sclerite, parameres short, curved, and pointed. 


12. Monohelea (Monohelea) nebulosa (Coquillett), 1901 
Ficurss 18, f; 19, ¢ 


Ceratopogon nebulosus Coquillett, Proc. U. 8. Nat. Mus., vol. 23, p. 606, 1901 
(os, New Jersey). 

Ceratolophus nebulosus, Kieffer, Genera insectorum, fasc. 42, p. 60, 1906. 

Johannseniella nebulosa, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, 
p. 226, 1914. 

Hartomyia nebulosa, Malloch, Bull. Illinois State Lab. Nat. Hist., vol. 10, p. 340, 
1915 (Indiana). 

Monohelea nebulosa, Kieffer, Ann. Mus. Nat. Hungarici, vol. 15, p. 312, 1917.— 
Johannsen, Ann. Ent. Soc. Amer., vol. 36, p. 781, 1943 (Massachusetts, 
Idaho). 

Description.—Femate: Length 2.1 mm., wing 1.7mm. by 0.6 mm. 
Head dull, dark brown, with a few pruinose gray spots on vertex; 
antennae light brown, distal segments not darker. Palpi light brown. 

Mesonotum pruinose gray, with scattered, small, dark-brown dots, 
each mesonotal hair arising from a dot, these dots more or less con- 
fluent, especially on posterior portion of mesonotum. Scutellum 
brown, middle third yellow, with about eight marginal bristles. 
Postscutellum and pleura dark brown. Legs shining dark brown, 
knees and tarsi yellowish. Posterior femora and tibiae greatly 
thickened; hind basitarsus with basal and distal spine; second segment 
with two distal spines, third segment with one distal spine; claws 
minute and equal on forelegs and midlegs, outer claw very long, 
four times the length of inner, on hind legs. Proportions of segments 
of hind leg as in table 1. 

Wing grayish hyaline, a quadrate black patch across first basal cell 
at half its length; two broad, irregular, more or less interconnected, 
transverse bands across wing, the first at first radial cell, the second at 
apex of second; cell R; thus with a dark, a light, a dark and a light band, 
all of subequal breadth. First radial cell 0.4 times as long as second. 
Halteres pale yellow. 

Abdomen uniform, pruinose, dark brown; anal segment yellow. 

Mate: Ninth sternite transverse, without caudal lobe, posterior 
portion spiculate; ninth tergite very narrow, with sides subparallel on 
distal half, apicolateral processes short and rounded. Basistyles very 
stout and convex; dististyles slender and curved, 0.6 as long as basi- 
styles. Aedeagus with two sections, basal part about as broad as long, 
anterior margin contiguous with sternite, posterior margin deeply 
emarginate, the slender, rodlike, distal sclerite with notched apex fit- 


152 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


ting into the emargination. Parameres connected by a transverse 
bridge near bases, with slender anterior and lateral apodemes; each 
paramere with posterior portion narrowed and curved ventrolaterad 
to a bladelike apex, a small sub-basal tooth on ventral side. 

Material eramined.—New York: Farmingdale, July 9, 1938, H. and 
M. Townes, 1 o*; Millwood, June 21, 1936, H. Townes, 1 9. 

New Jersey: Atsion, June 17, 1949, H. Townes, 1 9; Medford Lakes, 
June 17, 1939, H. Townes, 1 o; Riverside, June 18, 1939, H. Townes, 
1 9; Riverton, June 19, Johnson, 1 & (type). 

Georgia: Thomasville, May 15-30, Palmer, 1 9. 

Arkansas: Pike County, June 12, 1938, Turner, 1 9. 

Remarks.—This species is very similar to the Palearctic species M/. 
(M.) tessellata (Zetterstedt), 1850, which, however, has on the scutel- 
lum four marginal bristles, the ends of which are yellowish, as is the 
median area. The wing markings of tessellata are much more nearly as 
in M. (M.) johannsent, new species. 

A male of tessellata from Norfolk, Hatfield, England, was examined 
through the kindness of Paul Freeman of the British Museum (Natural 
History). The specimen is unsatisfactory for a description of external 
characters, as it was glued to a card when wet. It is noted, however, 
that both the first and second segments of the hind tarsi bear a distal 
pair of spines. The aedeagus of the genitalia consists of a heavily 
sclerotized basal plate about three times as broad as long, bearing a 
small, crescentic, median sclerite on the ventroposterior margin and 
bearing dorsoposteriorly a large, triangular structure with sharp- 
pointed, median apex, and widely flaring, lateral arms articulating with 
inner margins of basistyles. Parameres with slender anterior arms 
connected by a broad, median bridge, the posterior portions simple, 
the basal halves stout, the distal halves very slender and curved ventro- 
cephalad. 

According to Tokunaga’s (1940) description, his record of tessellata 
from Japan probably refers to an as yet unnamed species. This Jap- 
anese species has the scutellum yellow only in the middle, as in nebulosa, 
which it also closely resembles in wing markings; but the mesonotum 
has distinct yellow spots, the antennae have the distal segments and 
the apices of the basal segments brownish, the legs are yellowish brown 
with dark brown markings, and the dorsum of the abdomen is pale. 
yellowish brown. 

The West African species M. (M.) litoraurea Ingram and Macfie, 
1921, (female) and M. (M.) mimas de Meillon, 1939, (male) are very 
similar to M. (M.) nebulosa (Coquillett). The genitalia of mamas are 
the same as those of nebulosa except that the dististyles are abruptly 
bent near the base. 


HELEID MIDGES, GENUS MONOHELEA—WIRTH 153 


13. Monohelea (Monohelea) johannseni, new species 
Figures 18, g, k, 1; 19, a 


Monohelea tessellata, Johannsen, not Zetterstedt (misidentification), Ann. Ent. 
Soc. Amer., vol. 36, p. 781, 1943 (Alabama). 

Diagnostic characters——A dark brown species with large pruinose 
gray mesonotal markings, black scutellum and legs, and maculate 
wings. 

Description —Frmate: Length 1.5 mm., wing 1.2 mm. by 0.5 mm. 
Closely resembling MZ. (1Z.) nebulosa (Coquillett) but smaller, the 
mesonotum dark brown with large, irregular, pruinose gray areas, the 
scutellum entirely blackish and with four long and several shorter 
marginal hairs. Proportions of hind leg of female as in table 1, hind 
tarsi with one basal and two distal spines on first segment, two spines 
at tip of second and occassionally two spines at apices of third and 
fourth segments; inner claw on hind leg of female about a third as 
long as outer. Wing as in nebulosa, but the markings behind vein 
M, much fainter than those on anterior part of wing. Spermathecae 
two, subequal, oval, each with very short, sclerotized duct. 

Male genitalia as in nebulosa, but the parameres simple, without 
the sub-basal ventral tooth, and distinctly and abruptly bent mesad 
at distal fifth. 

Types.—Holotype, 2, USNM 61095, Falls Church, Virginia, July 8, 
1950, Wirth. Allotype, Innerarity Point, Escambia County, Florida, 
Apr. 29, 1949, Rathert. Paratypes: Virginia: 2 99, Mountain Lake, 
July 15, 1938, July 21, 1940, L. and M. Milne. Florida: 4 #0, 5 99, 
same data as allotype except date April and May 1949; 1 9, Santa 
Rosa, Walton County, May 5, 1950, Peterson. Alabama: 1 9, La- 
Place, June 9, 1917, Bradley. Michigan: 1 9, Cheboygan County, 
July 17, 1942, Sabrosky. Iowa: 1 9, Pikes Peak State Park, Clayton 
County, July 4, 1949, Laffoon. 


References 


Fiocs, H., and ABONNENC, E. 

1942. Cératopogonidés divers de la Guyane Francaise III. Inst. Pasteur 

Guyane Terr. L’Inini Publ. 55, 6 pp., 2 figs. 
GoETGHEBUER, M. 

1934. Heleidae (Ceratopogonidae), Jn Lindner, Die Fliegen der Palaearkti- 

schen Region, Lief. 78, pp. 49-94. 
JOHANNSEN, O. A. 

1943. A generic synopsis of the Ceratopogonidae (Heleidae) of the Americas, 
a bibliography, and a list of the North American species. Ann. 
Ent. Soc. Amer., vol. 36; pp. 763-791. 

KIEFFER, J. J. 

1917. Chironomides d’Amérique conservés au Musée National Hongrois de 

Budapest. Ann. Mus. Nat. Hungarici, vol. 15, pp. 292-364, 43 figs. 
LANE, J. 

1945. Redescrigéo de Ceratopogonideos Neotrépicos (Diptera: Ceratopogo- 

nidae). Rev. Ent., vol. 16, pp. 357-372, 19 figs. 
Leg, D. J. 

1948. Australasian Ceratopogonidae (Diptera, Nematocera). Part 4. The 
Stilobezzia group of genera. Proc. Linn. Soc. New South Wales, 
vol. 72, pp. 345-356, 1 pl., 23 figs. 

Macris, J. W.S. 

1937. Ceratopogonidae from Trinidad. Ann. Mag. Nat. Hist., ser. 10, vol. 
20, pp. 1-18, 6 figs. 

1940a. A report on a collection of Ceratopogonidae (Diptera) from British 

Guiana. Ent. Monthly Mag., vol. 76, pp. 23-32, 4 figs. 
1940b. Ceratopogonidae (Diptera) from British Guiana and Trinidad. 
Proc. Ent. Soc. London, ser. B, vol. 9, pp. 179-195, 4 figs. 
Toxunaaa, M. 

1940. Chironomoidea from Japan (Diptera), XII. New or little-known 
Ceratopogonidae and Chironomidae. Philippine Journ. Sci., vol. 
72, pp. 255-311, 4 pl. 

WirrTu, W. W. 

1952. The Heleidae of California. Univ. California Publ. Ent., vol. 9, pp. 
95-266, 33 figs. 

154 


U. S. GOVERNMENT PRINTING OFFICE: 1953 


PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


issued "4x 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington : 1953 No. 3321 


A REVIEW OF THE BEETLE FAMILY CEPHALOIDAE 
By Ross H. Arnett, Jr.’ 


The family Cephaloidae consists of only one genus, Cephaloon 
Newman, 1838, with eight known species that are locally rather com- 
mon but are known only from the Eastern United States, Western 
United States, Western Canada, Japan, and Amur. All of the species 
are closely related and quite variable in color. Useful separation 
characters are to be found in the antennae and various male structures. 
As yet little is known concerning the habits and life histories of mem- 
bers of this group. All known species of this family are represented 
in the collections of the U. S. National Museum. 

After study of some material from the Ussuri River Valley recently 
sent to me for determination, it seems desirable to reconsider the 
taxonomy, affinities, and distribution of this group. The ranking of 
this small assemblage of species as a family on equal ground with the 
rest of the families in the order is still somewhat doubtful in my mind. 
I believe the placing of the Cephaloidae as a satellite of the great 
family Tenebrionidae is firmly established, but my studies in the 
Tenebrionoidea have not as yet revealed, on the basis of our present 
ranking and evaluation of what defines a family, any group with which 
the genus Cephaloon can be incorporated; hence, I retain it as a family. 

I wish to thank Mr. Hugh B. Leech and Dr. E. C. Van Dyke, both 
of the California Academy of Sciences, and Dr. Hans Klapperich of 
Bonn, Germany, for the loan of several specimens used in this study. 


1 Bureau of Entomology and Plant Quarantine: U. S. Department of Agriculture. 
232728—53 155 


156 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Family CepHALOIDAE LeConte 


Cephaloidae LeConte, Smithsonian Misc. Coll., vol. 3, art. 3, Classification of the 
Coleoptera of North America, pt. 1, p. 259, 1862. 


LeConte originally proposed this family for the North American 
Cephaloon lepturides Newman, but also referred to two species, al- 
though not by name, which had been described by Motschulsky from 
the Amur River Valley. Several genera have since been erected for 
species in this group, and then synonymized, until in the present 
study the family contains only the original genus Cephaloon Newman. 

Family diagnosis —Size 8-20 mm.; head elongate, diamond-shaped, 
deflexed; antennae 11-segmented, filiform, with apical segments some- 
what enlarged, inserted between the eyes and bases of the mandibles; 
mandibles elongate, acute at apex, never bifid or otherwise modified; 
eyes reniform, not prominent; maxillary palpus 4-segmented, first 
segment small, obscure, apical segment triangular. Pronotum with- 
out lateral margins, smooth, always abruptly narrowed anteriorly 
from the middle. Legs slender; front coxal cavities open behind; 
front and middle coxae prominent, conical; apical spurs of all tibiae 
large, two on each tibia; tarsi 5-5-4, the segments all simple, not lobed 
or tomentose beneath; claws pectinate, with a subequal membranous 
lobe beneath each claw. Elytra with vague costae, minutely punctate, 
never striate. Abdomen with 5 visible sternites in the female, 7 in 
the male (sternites 2 +3 to 9 visible inthe male). Body covered with 
very fine pubescence. 

MALeE GENITALIA: Apical abdominal segments (7-9) of the male 
considerably modified. Segment 7 with sternite and tergite laterally 
fused, forming a globular shaped segment. Segment 8 with sternite 
and tergite laterally fused, forming two laterally triangular pieces. 
Segment 9 closely fitted into segment 8, the tergite and sternite both 
triangular and filling the emarginations of segment 8. Genital organs 
themselves quite simple; paramere two short, freely articulate lobes 
fitted onto the apex of the large troughlike basal piece which more or 
less envelops a simple, small, somewhat curved, and tubelike median 
lobe; no evidence of a tegminite. 

Affinities of the family—The heteromerous tarsi place this family 
in the Tenebrionoidea; the open anterior coxal cavities place it*in the 
group including the Oedemeridae, Pythidae, and Serropalpidae, as 
distinguished from the group including the Alleculidae,* Lagriidae, 
and Tenebrionidae, all of which have closed coxal cavities. The ser- 
rate claws with fleshy pulvilli and the laterally fused eighth abdominal 
segment of the male separate it from the others of this group of Teneb- 
rionoidea. The very small median lobe and the larger paramere 
with the small lateral lobes show affinities with the genus Mycterus of 


BEETLE FAMILY CEPHALOIDAE—ARNETT 157 


ce 


Wes ~ 


© . lepturides Newm. . 
C. bicolor Horn 
& C. tenuicome LeC. ane 
@® C. vardykei Hopp. & Hopp. | 5 ee 
Ria ee 


A C. pacificum VanD. 
B 


OC. ungulare LeC. 


Y C. pallens Mots. 
® C. variabilis Mots. 


Ficure 20.—Distribution of the family Cephaloidae. 


158 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


the family Pythidae. In addition, Mycterus and Cephaloon both 
have prominent procoxae and mesocoxae and the prothorax is with- 
out lateral margins. Most workers in the past have considered the 
Oedemeridae and Cephaloidae closely allied, but my incomplete 
studies of the male genitalia of the heteromerous beetles indicate that 
this is not so. The very different structure of the apical abdominal 
segments of the species of Cephaloon leads me to suppose that these 
species are far removed from the oedemerids, but their true affinities 
are not yet recognized. 

Notes on the distribution of the species—As can be seen from the 
accompanying map (fig. 20), the species of this family show the typical 
distribution pattern of a Holarctic group which was probably derived 
from some Asian stock and spread to North America via the Alaskan 
land bridge. It is interesting to note that, based on both external 
morphology and the morphology of the male genitalia (in the case 
of C. pallens) both of the Asian species are most similar to two of the 
Eastern North American species and not to the Western North 
American species. If these similarities of morphology reflect relation- 
ship, as we assume they do, then we again see the often-repeated 
pattern of a pre-glacial distribution across northern Canada from 
Alaska, down through Ontario and into Eastern United States. 
This I believe to be a further stock-piling of evidence against the 
theories of continental drift and the North Atlantic land bridge idea. 


Genus Cephaloon Newman 


Ichnodes Dejean, 1834, Cat. Col. ed. 3, p. 227. (Nomen nudum, one ms. trivial 
name listed.) 

Cephaloon Newman, Ent. Mag., vol. 4, p. 376, 1838. (Genotype, Cephaloon 
lepturides Newman; 1838; monobasic.) 

Cephalaon Motschulsky in Schrenck, Reisen und Forschungen in Amurlande, 
vol. 2, pt. 2, p. 140, 1860. (Error for Cephaloon.) 

Typitium Casey, Ent. News, vol. 9, p. 193, 1898. (Genotype, Cephaloon ungulare 
LeConte, Proc. Boston Soc. Nat. Hist., vol. 16, p. 275, 1873; original desig- 
nation and monobasic. Placed in synonymy by Hopping and Hopping, 
Pan-Pacific Ent., vol. 10, p. 64, 1934.) 

Sponidium Casey, Ent. News, vol. 8, p. 193, 1898. (Genotype, Cephaloon 
tenuicorne LeConte, 1873; present designation. Placed in synonymy by 
Hopping and Hopping, 1934.) 

Ephamillus Semenow, Horae Soc. Ent. Ross., vol. 34, p. 495, 1900. (New 
synonymy. Genotype, Cephaloon variabile Semenow =Cephaloon variabilis 
Motschulsky in Schrenk, 1860; original designation and monobasic.) 

Drachylis Casey, 1898, Ent. News, vol. 9, p. 195. (New synonymy. Genotype, 
Drachylis simulans Casey, 1898; monobasic.) 


Discussion—The type and only known specimen of Drachylis 


simulans Casey, the genotype of Drachylis Casey, has been examined. 
This is an unfortunate example of the description of a species on one 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 5 


Adult, male genitalia, and claws of Cephaloidae: 1, Cephaloon itenuicorne LeConte, female, 
showing general habitus; 2, paramere of male, C. tenwicorne; 3, paramere of male, C. 
bicolor Horn; 4, paramere of male, C. pacificum Van Dyke; 5, paramere of male, C. 
lepturides Newman; 6, apical portion of paramere of male, C. ungulare LeConte; 7 
abdominal segments 7-9 of male, C. ungulare; 8, median lobe of male, C. pallens 
Motschulsky; 9, median lobe of male, C. lepturides Newman; 10, hind claw of C. tenui- 
corne; 11, hind claw of C. pallens; 12, hind claw of C. variabilis Motschulsky. 


-“<— 


BEETLE FAMILY CEPHALOIDAE—ARNETT 159 


poor specimen. Casey was certainly justified in wanting to describe 
this specimen for he believed that it lacked the comblike claws charac- 
teristic of the other species in the family, and therefore, its description, 
even though based on a single specimen, would alert collectors to hunt 
for such astrange cephaloid. <A close examination, however, reveals it 
to have been patched, the legs being undoubtedly from some other 
beetle. It appears to be Cephaloon bicolor Horn, with which I synony- 
mize it. 

The genus Ephamillus Semenow is based,on the same variable 
characters possessed by the other genera erected for species of this 
family. Kéno (Fauna Nipponica, vol. 10, fasc. 8, No. 10, pp. 76-82, 
1937) illustrates three characters, which, if they were constant and as 
illustrated, would serve for recognition of a genus. However, none of 
them appears to be constant or as distinctive as thought by Kéno and 
others. The acute pulvilli, curved at the tips, are found in three 
species, C. pacificum, C. ungulare, and C. variabilis. The shape of the 
pronotum, as illustrated by Kéno for C. variabilis, is subject to the 
same sort of variation in all the species. Finally, the sinuation of the 
hind tibia of C. variabilis, reported to be so pronounced that a portion 
of the tibia is thrown out of line at least a distance equal to the width 
of the tibia, is often barely perceptible even under a microscope, and 
on some specimens of the species it cannot be seen at all. For these 
reasons, I feel that this genus is invalid and I place it in synonymy 
with Cephaloon. 

The marginate pronotum eliminates the genus Stenocephaloon Pic, 
1932 (Mélanges Exotico-Entomologiques, fasc. 59, p. 2; genotype, 
Stenocephaloon metallicum Pic, monobasic) from this family. Until 
specimens can be studied, it is best placed in the family Serropalpidae 
near the genus Stenotrachelus Berthold. 

The following key to the known species of this family is adapted 
from Hopping and Hopping (Pan-Pacific Ent., vol. 10, pp. 64-70, 
1934). For identification purposes, the illustrations which accompany 
that paper are very useful. 


Key to the species of Cephaloidae 


1. Pulvilli of tarsal claws slender, acute, and curved at tips (pl. 5, fig. 12)____- 2 
Pulvilli of tarsal claws robust, obtuse, not curved at tips (pl. 5, figs.10, 11). 4 

2. Three distal antennal segments together approximately 3 mm. long; hind 
femora at most simply curved; 10 to 15 mm. (United States) _-_-------- 3 
Three distal antennal segments together not over 1.5 mm. long; hind femora 
tending to be sinuate; size 17 to 20 mm. (Amur River, Eastern Siberia, and 
Papa) osveenus.» 25. hohhse od hizde see C. variabilis Motschulsky 


160 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


3. Lateral margins of pronotum behind middle distinctly femarginate; Western 
(Uintted Stateses0-).2 bo tap er ee C. pacificum Van Dyke 
Lateral margins of pronotum behind middle almost straight (Eastern United 
is, {C7 5) plein ie pa ei et ches 2.6 os ordre meant ei A 5. C. ungulare LeConte 
4, Antennae with last three segments thickened___________-_._____-_____ 5 
Antennae with last three segments not markedly thickened____________- 6 

5. Elytra without sutural and marginal black stripes (Eastern United States). 
C. lepturides Newman 
Elytra with sutural and marginal black stripes (Amur River, Eastern Siberia 
CNLOC 0 LIPL ES) cy 8 Oe Ac eel Ac Re a A at hc C. pallens Motschulsky 
6. Three distal antennal segments together approximately 1.5 mm. long (West 
Coastiof Wmited-Stathans 2 iicsl. evil Bel 98 eee C. bicolor Horn 
Three distal antennal segments together barely over 1 mm. long (Western 
United, States andCanada)= - 2.242222). -2. 82.2.2. 7 
7. Male with bifurcate process of third visible abdominal sternite long and finger- 
like; female with apical abdominal sternite shallowly and narrowly emargi- 
nate; color of female black to testaceous___________ C. tenuicorne LeConte 
Male with bifurcate process of third visible abdominal sternite shorter, more 
triangular; female with apical abdominal sternite deeply emarginate; color 

of female reddish testaceous with elytra and metasternum black. 

C. vandykei Hopping and Hopping 


Key to the species of Cephaloon, based upon the male genitalia 


Male of Cephaloon variabilis Motschulsky unknown. 
1. Ratio of length of paramere to length of basal piece not over 1:2________ 2 
Ratio of length of paramere to length of basal piece not less than 1:3.5____ 3 
2. Median lobe tapering abruptly at apical third, slender (pl. 5, fig. 8); paramere 
tobes long And) slender. 2005. SC C. pallens Motschulsky 
Median lobe evenly tapering from base, more robust (pl. 5, fig. 9); paramere 
lobes shorter and heavier (pl. 5, fig. 5)-...____.-_~- C. lepturides Newman 
3. Paramere lobes acute at apex (pl. 5, fig. 6); median lobe long and slender, 
BIGGS MOLe PAPAUCL 452k Se cee s Se ee ee eee C. ungulare LeConte 
Paramere lobes enlarged at apex; median lobe shorter and more robust at 
QPCXas 2. Ss Secs Sec Sess soe ES ee 4 
4. Apex of paramere lobes about twice width of base when viewed laterally 
Klemen gll oes) ae ee C. vandykei Hopping and Hopping and C. bicolor Horn 
Apex of paramere lobes about same width as at base_________________-_ 5 
5. Apex of basal piece laterally extending beyond base of paramere to about 
one-half length of paramere lobes; paramere lobes stout, blunt at apex, 
uniform, in “width (pl: b; fig, 4) 2022. 2222 ee C. pacificum VanDyke 
Apex of basal piece laterally extending only slightly beyond base of paramere; 
paramere lobes more slender, and slightly enlarged at apex (pl. 5, fig. 2). 
C. tenuicorne LeConte 


In view of the relatively recent and thorough discussion of the 
species by Hopping and Hopping (1934), I feel that it is superfluous 
to repeat it here except to mention that under C. bicolor the name 
Drachylis simulans Casey should be added as a synonym (see discus- 
sion under generic synonymy preceding). 


BEETLE FAMILY CEPHALOIDAE—ARNETT 161 


There follows the bibliographic citations of the two exotic species 
not mentioned by Hopping and Hopping. The key and illustrations, 
I believe, sufficiently characterize them. A revised section of the 
Coleopterorum Catalogus (Junk) is being prepared which will give all 
of the literature references, therefore they will not be cited here. 


Cephaloon pallens Motschulsky, 1860 
PuaTE 5, Figures 8, 11 


Cephaloon pallens Motschulsky, in Schrenck, Reisen und Forschungen in Amur- 
lande, vol. 2, pt. 2, p. 140, pl. 9, fig. 15, 1860. (Type locality, Kisi, on the 
River Amur.) 

Cephaloon pallens var. cinctipennie Heyden, Deutsche Ent. Zeitschr., p. 167. 
1892. (Type locality unknown, except ‘‘Amur”’ as given in the title of the 
paper.) 

Cephaloon pallens var. koltzei Heyden, Deutsche Ent. Zeitschr., p. 168, 1892. 
(Type locality, ‘‘Amur,”’ as above.) 

Cephaloon pallens var. maculicolle Heyden, Deutsche Ent. Zeitschr., p. 167, 1892. 
(Type locality, ‘‘Amur,” as above.) 

Cephaloon pallens var. picticolle Heyden, Deutsche Ent. Zeitschr., p. 167, 1892. 
(Type locality, ‘‘Amur,” as above.) 

?Cephaloon pallens Motschulsky, Solsky, Horae Soc. Ent. Rossicae, vol. 11, p. 
295, 1875. (Djalinda River. Note: This is probably a misidentification of 
Ephamillus variabilis var. tristiculus Heyden, 1892, judging from the descrip- 
tion and locality.) 

Cephaloon sakurae Lewis, Ann. Mag. Nat. Hist., ser. 6, vol. 15, p. 444, fig. 10, 
1895. (Placed as synonym by Kéno, 1937, Fauna Nipponica, vol. 10, fasc. 
8, No. 10, p. 79.) 


In a species with the extreme color variation exhibited in all of the 
species of this family, the naming of a few of the color variants serves 
no useful purpose and these are therefore here disregarded. 


Cephaloon variabilis Motschulsky, 1860 
PuaTE 5, FigurE 12 


Cephaloon variabilis Motschulsky, in Schrenck, Reisen und Forschungen in Amur- 
lande, vol. 2, pt. 2, p. 141, 1860. (Type locality, Mariinsk on the River 
Amur.) 

Cephaloon variabilis var. tristiculus Heyden, Deutsche Ent. Zeitschr., 1892, p. 169. 
(Type locality, unknown except ‘‘Amur.’’) 


Here again, the naming of a single color form can serve no useful 
purpose. 


WW. S. GOVERNMENT PRINTING OFFICE: 1953 


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SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington : 1953 No. 3322 
THE FRESH-WATER TRICLADS (TURBELLARIA) OF ALASKA 


By Roman KENnK 


Introduction 


In 1948, during an investigation of biting insects in Alaska con- 
ducted by the Bureau of Entomology and Plant Quarantine, United 
States Department of Agriculture, Dr. Reese I. Sailer collected and 
transmitted to me several samples of fresh-water triclads (planarians). 
An examination of the material revealed that the worms belonged to 
the genus Polycelis, a genus fairly common in Asia and Europe but 
only twice reported from North America. The finding suggested a 
closer relationship of the Alaskan fresh-water fauna with that of East 
Asia. A more thorough study of the triclads of Alaska promised to 
yield a more definite understanding of these zoogeographical relations. 

My field trip 1 to Alaska was made in the summer of 1950. Since 
the available time was rather limited, the collecting was done mainly 
along the highways of the territory and in the vicinities of Point 
Barrow and Umiat. 

The following roads and fresh-water localities were visited on the 
trip: Steese Highway in the section between Fairbanks and Birch 
Creek (milepost 103); Elliot Highway, from Fairbanks to Livengood; 
tundra lakes and pools in the vicinity of Point Barrow; Colville 
River, several streams, and a lake near Umiat; Mount McKinley 
Park Road, a section of about 15 miles adjoining the railroad station; 
Glenn Highway, from Anchorage to Glenallen; road from Palmer to 

1 This study was supported by the Arctic Institute of North America under contractual arrangement 
with the Office of Naval Research. 
238538—53——1 163 


164 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Willow; road from Anchorage to Potter; and Richardson Highway, 
section between Valdez and Glenallen. 

No fresh-water triclads were found by me either at Point Barrow 
or at Umiat. I received, however, from the U.S. National Museum, 
a sample of triclads collected by P. F. Scholander in a lake near Umiat. 
The relative scarcity of planarians in tundra lakes and pools may be 
due to the high acidity of the waters or to the extremely severe con- 
ditions prevailing in them during the long winter season. 

Grateful acknowledgment is made of the very helpful cooperation 
which was extended to me by several agencies in Alaska: The Arctic 
Research Laboratory, Point Barrow; the U.S. Public Health Service, 
Anchorage; the Alaska Road Commission, Fairbanks, Anchorage, and 
Glenallen; and the Division of Forestry, Bureau of Land Management, 
Fairbanks and Anchorage. I also wish to express my indebtedness to 
Prof. Edward G. Reinhard of the Catholic University of America and 
to Dr. Fenner A. Chace, Jr., and Dr. Doris M. Cochran of the U. S. 
National Museum who kindly permitted me to use their laboratory 
and office facilities in Washington, D. C. 

Four species of fresh-water triclads were collected in Alaska. Two 
are inhabitants of the White Mountains, a mountain range extending 
in an east-west direction north of Fairbanks. The other two are 
widely distributed in waters of the Alaska Range and of the southern 
section of Alaska and one of them reaches as far north as Umiat. 


Family PLANARIIDAE 
Genus Phagocata Leidy 


Phagocata nivea, new species 
Figure 21; Puate 6, Figure 1 


Description.—This is a slender, rather delicate species. Mature 
specimens measure up to 8 mm. in length and about 1.5 mm. in width. 
In the quietly gliding animal the anterior end is truncated,with a very 
slightly bulging frontal outline and with rounded lateral corners 
(auricles). ‘There is no distinct narrowing or neck behind the auricles 
and the lateral margins of the head are approximately parallel. Behind 
the head, the body widens and soon reaches its greatest width. From 
there on, the lateral margins of the body run parallel up to the level 
of the mouth, to converge again in the postpharyngeal region and to 
meet in a bluntly pointed posterior end. 

The species lacks pigment and usually appears white and somewhat 
transparent. The intestinal contents may shine through the body 
wall and give the animal a certain amount of color; the margins of the 
body, the head region, and the areas occupied by the pharynx and the 
copulatory apparatus, however, are always white. 


FRESH-WATER TRICLADS OF ALASKA—KENK 165 


There are two rather small eyes, situated close together (about one- 
fourth the body width apart at the level of the eyes) and far removed 
from the frontal end. This character, easily recognized in life, dis- 
tinguishes the species from another white triclad with which it shares 
its habitat, Dendrocoelopsis alaskensis, described as new on p. 178. 

The pharynx is inserted, in sexually mature specimens, somewhat 
behind the middle of the body and measures about one-sixth of the 
body length. The copulatory organs occupy the anterior half of the 
postpharyngeal region. 

The animal moves by gliding only; crawling movements, such as 
are seen in other triclads, particularly in those equipped with anterior 
adhesive organs, have not been observed in this species. 

From the description it may be seen that the species in life shows 
a close resemblance to other species of the same genus, particularly 
to the American Phagocata morgani, the European P. albissima, P. 
vitta, and related forms. A separation of these species can be made 
only on the basis of anatomical characters. 

Only those characters of the digestive system that have a taxonomic 
significance are discussed here. The pharynx has a structure typical 
of the family Planariidae; i. e., the fibers of the internal muscle zone 
are arranged in two distinct layers, a thick inner circular layer and a 
narrower outer longitudinal one. The anterior intestinal trunk bears 
10 or 11 branches on each side. Each posterior trunk has 21 to 27 
lateral branches and numerous short medial branches in both the 
pharyngeal and postpharyngeal regions. 

The testes are numerous and are arranged, on each side of the 
midline, in a longitudinal zone extending from a short distance behind 
the ovary almost to the posterior end of the body. Their position is 
predominantly ventral, below the intestinal branches. Only a few 
testes extend into the mesenchymatic ‘‘septa’’ between the branches 
toward the dorsal side. 

The two ovaries are typical, each situated approximately below the 
second intestinal branch. An undifferentiated mass of cells, the par- 
ovarium, is attached to the dorsolateral side of each ovary. 

The genital pore (pg), situated about halfway between the mouth 
and the posterior end of the body, leads into a small cavity (ac) 
which continues to the left and dorsally into the duct of the copulatory 
bursa (bd) and to the right and anteriorly into the male atrium (am). 
This cavity may be considered to represent a common genital atrium. 
In some specimens, however, there appears to be no differentiation 
of the atrium into male and common parts, and the bursa duct and 
an undivided atrium meet at or near the genital pore. These varia- 
tions are obviously due to the different states of muscular contraction 
in which the animals were killed. The atrium, narrow at the genital 
aperture, widens as it extends forward, to the right side of the midline. 


166 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


It is lined with a tall, glandular epithelium, the cells of which project 
into the cavity in a villuslike fashion. Below the epithelium there 
are two layers of muscle fibers, one circular and the other longitudinal. 

The penis consists of a spherical, muscular bulb embedded in the 
mesenchyme, and a moderately large papilla projecting into the male 
atrium. The bulb is pierced by numerous gland ducts which open 
into the lumen of both the bulb and the papilla. The shape of the 
papilla is subject to great variation, due apparently to the state of 
contraction of the organ. It may be twisted to one side and even 
partly inverted into the lumen of the penis (similar to the pseudo- 
flagellum of various dendrocoelids). The shape shown in figure 21 
appears to be that of the organ at rest. The outer wall of the papilla 
is covered with a tall to cubical epithelium similar to that lining the 
atrium. Below the epithelium there is a layer of circular muscle 
fibers followed by another of longitudinal fibers. The shape of the 
penis lumen (/p) is as changeable as that of the papilla. Typically, 
it appears to be wider in the bulb than it is in the papilla, though 
there is no distinct ejaculatory duct differentiated. The lumen opens 
ventrally to the tip of the papilla. The two vasa deferentia (vd) 
penetrate the penis bulb from both sides and empty into the penis 
lumen separately, but not far apart. 

The two oviducts converge at the level of the copulatory apparatus, 
the left one passing between the bursa duct and the male atrium, and 
unite at a point dorsally to the atrium. The rather long common 
oviduct (odc) curves ventrally and opens into the posterior part of the 
male atrium. The terminal sections of the paired oviducts and the 
greater part of the common oviduct receive the outlets of numerous 
eosinophilic glands, the cell bodies of which are scattered in the sur- 
rounding mesenchyme, particularly dorsally to the atrium. 

The copulatory bursa (6) is of moderate to large size and is ir- 
regularly lobed. The bursa duct or stalk (6d) is wide, runs posteriorly 
to the left of the midline, and curves ventrally to reach the genital 
aperture. It is lined with a tall, glandular epithelium and surrounded 
with a strong muscular coat consisting of mtermingled circular and 
longitudinal fibers. 

Taxonomic position—The genus Phagocata Leidy (Fonticola 
Komarek) in its present extent (cf. Hyman, 1937, pp. 300-302) has 
representatives in Europe, Asia, and North America. The genus is 
not quite homogeneous and will probably, in due time, be subdivided 
into several genera (cf. Beauchamp, 1939). For the purpose of com- 
parison, we may consider here only those species of the genus that 
lack pigment. Though the presence, or the lack, of pigment is a 
character of subordinate taxonomic value, it may nevertheless serve 
well as a character of specific rank. The Alaskan form differs from 


FRESH-WATER TRICLADS OF ALASKA—KENK 167 


European species such as P. vitta (Dugés) and P. albissima (Vejdovsky) 
and from the American white P. morgani (Stevens and Boring) mainly 
in the structure of the male copulatory organ. Asia has several 
species of Phagocata, the majority of them pigmented forms. Of the 
three unpigmented Asiatic species that may belong to the genus, two 


Figure 21.—Phagocata nivea, diagram ot the copulatory organs in longitudinal section, 
X 92. ac, common atrium; am, male atrium; b, copulatory bursa; bd, bursa 
stalk; /p, penis lumen; 0, mouth; odc, common oviduct; pg, genital pore; 2d, vas deferens. 


have been described from immature specimens and the anatomy of 
their reproductive systems is not known: Planaria pellucida Yjima 
and Kaburaki (1916) from Sakhalin and a species from the Baikal 
region assigned tentatively to Fonticola by Bazikalova (1947). <A 
third species, Phagocata coarctata (Arndt, 1922), from the vicinity of 
Vladivostok, is sufficiently well known, although no fully mature 
individuals of this species have been studied. P. coarctata differs 
from P. nivea externally in being smaller and broader, and in having 
a different contour of the anterior end, which bears protruding lateral 
lobes, and a greater distance between the two eyes. Anatomically, 
the two species are, undoubtedly, closely related.” 


2 Livanov and Zabusova (1940, p. 146) state that a reexamination of Arndt’s slides of Planaria coarctata 
showed that the arrangement of the muscle fibers of the pharynx conformed with the dendrocoelid type 
(circular and longitudinal fibers of the internal muscle zone intermingled). Arndt (1922, p. 108) described 
the anatomy of the pharynx in minute detail and indicated, both in a figure (pl. 4, fig. 7) and in the text, 
a typical planariid pattern. I must assume that some confusion occurred somewhere, probably in the 
identification of the slides sent to Livanov. 


168 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


In Phagocata nivea the penis lumen opens usually below the tip of 
the penis papilla. This character has been used by Livanov and 
Zabusova (1940, p. 96) to segregate a group of Asiatic species from 
Phagocata and to place them in a new genus, Penecurva. The same 
character is found, however, in a common North American species, 
Phagocata morgani, which shows no other close relations with the 
Asiatic group. This character is apparently inadequate as a basis 
for the establishment of a new genus. 

Holotype.—On one slide, USNM 22332, creek crossing Elliot High- 
way at milepost * 31.0, July 24, 1950. 

Distribution and ecology.—Phagocata nivea was collected in cool, 
fast mountain streams on the slopes of the White Mountains, a range 
north of Fairbanks. The temperature of the water ranged from 3.2 
to 7.2° C. (July). The animals are cold-stenothermic and do not 
tolerate sudden increases in temperature. They were often found 
in the company of another white triclad, Dendrocoelopsis alaskensis. 


Stream on Steese Highway (pl. 8), at milepost 84.0, altitude 2,700 feet, July 
19 and 21, 1950, water temperature 6.9° C.; one immature and one mature speci- 
men, from under stones. 

Willow Creek, on Steese Highway, at milepost 96.6, altitude 2,100 feet, July 
19, 1950, 7.2° C.; two immature specimens, from under stones. 

Spring and stream on Steese Highway, milepost 82.5, near Alaska Road Com- 
mission camp, July 21, 1950; two mature and six immature specimens, on the 
undersides of stones. 

Fox Gulch, on Elliot Highway, milepost 1.2, July 24, 1950, 5.0° C., one small 
specimen near bait (beef liver). 

Creek crossing Elliot Highway at milepost 31.0, July 24, 1950; 39 specimens, 
about half of them mature, collected under stones (holotype). 


Genus Polycelis Ehrenberg 
Polycelis borealis, new species 
FigurE 22; Puate 6, Figure 2 


Description —Mature animals are usually 12 to 15 mm. long and 
1.5 to 2 mm. wide (larger specimens, measuring up to 20 mm. in 
length, have been seen). The frontal margin is slightly convex. At 
the lateral corners of the head there is a pair of elongated, bluntly 
pointed auricles which are held raised when the animal is gliding 
quietly. Behind the auricles, the body first narrows slightly, then 
gradually widens, reaching its greatest width at the level of the 
pharynx. Behind the pharynx the body tapers to a moderately 
pointed posterior end. 

The color of the dorsal side is usually a uniform light or dark 
brown, that of the ventral side a light grayish brown. In animals 

3 The large highways of Alaska are marked with wooden mileposts indicating only full miles. Many 


posts were missing at the time when the collections were made. Fractional milages were usually estimated 
from the nearest milepost or from the speedometer readings of the car used. 


FRESH-WATER TRICLADS OF ALASKA—KENK 169 


from some localities, an indistinct lighter median line occurred dor- 
sally in the prepharyngeal region, with lighter areas above the pharynx 
and the copulatory organs. 

The species has many small eyes (a generic character) arranged 
in a band along the frontal margin of the head, the base of the auricles, 
and the lateral margins of the body a short distance behind the head. 
Anteriorly the eyes are placed in more than one row, somewhat 
irregularly scattered; behind the head they are in a single row reaching 
backward about one-third to one-half the length of the prepharyngeal 
region. There is no narrowing of the band of eyes at the base of the 
auricles. 

The pharynx is inserted at about the middle of the body and 
measures in length almost one-fourth the length of the body. The 
copulatory organs occupy more than half the post-pharyngeal region. 

The animal moves by gliding only. 

The pharynx of Polycelis borealis is structurally typical of the genus 
Polycelis and of the family Planariidae, the muscle zone being formed 
by two distinct layers, a thick circular layer adjoming the epithelium 
of the pharyngeal lumen and a thinner layer of longitudinal fibers. 
The anterior trunk of the intestine bears 5 to 6 lateral branches. 

The numerous, fairly large testes are arranged in two zones, to 
the right and left of the anterior intestinal trunk, extending from the 
level of the ovaries posteriorly to the base of the pharynx. The 
testes are essentially ventral, though individual vesticles may extend 
dorsally in the mesenchymatic spaces between the intestinal branches 
and occupy almost the entire dorsoventral diameter of the body. 
Where intestinal branches are present, however, the testes develop 
only below them. Each zone of testes reaches laterally only little 
beyond the ventral nerve cord. 

The ovaries are situated behind the first pair of lateral branches of 
the anterior trunk of the intestine. 

The genital pore (pg), situated in the midline behind the middle of 
the postpharyngeal region, connects with a narrow posterior extension 
of the genital atrium (a). This extension leads dorsally and somewhat 
to the left into the duct of the copulatory bursa and anteriorly into 
the widened portion of the atrium containing the penis. There is no 
marked division of the atrium into a posterior common atrium and 
an anterior male atrium. The wall of the atrium is lined with a 
cubical epithelium and equipped with two muscle layers, one circular 
and the other longitudinal. 

The penis consists of a bulb embedded in the mesenchyme and a 
papilla projecting into the atrium. Neither of the two parts is very 
muscular. The shape of the penis, particularly of the papilla, is very 


170 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


changeable. In specimens that were fixed in a well-extended condi- 
tion, the bulb appears spherical and the papilla has a conical shape. 
The bulb contains a spacious, irregularly lobed cavity, the seminal 
vesicle (vs). Numerous gland ducts empty into the lumen of the 
bulb after entering the bulb, particularly from its anterior surface, 
and penetrating between its muscle fibers. The ducts are filled 
with a granular, slightly eosinophilic secretion. The two vasa 
deferentia (vd) open into the seminal vesicle separately from the 
anterolateral sides. 

The penis papilla is conical or finger-shaped when well extended. It 
is covered with a cubical epithelium below which there are two rather 
feebly developed muscle layers, tapering in thickness toward the tip 
of the papilla: a layer of circular fibers adjoiming the epithelium and 
a thinner layer of longitudinal fibers. The absence of a strong mus- 
cular wall probably accounts for the great variation in the shape of 
the penis papilla observed in the material. The lumen of the seminal 
vesicle continues into the papilla as a wide canal (de), opening at or 
near (ventrally to) the tip of the papilla. The canal is lined with an 
epithelium of cubical cells and lacks the gland openings character- 
istic of the seminal vesicle. It corresponds to an ejaculatory duct 
but it apparently has no distinct muscle coat. 

The two oviducts, running posteriorly along the ventral nerve 
cords, bend dorsally and medially at the level of the copulatory 
organs. The left one passes through the space between the atrium 
and the bursa duct and unites with the oviduct of the right side 
dorsally to the atrium. The common oviduct (odc) runs ventrally 
and opens into the posterior part of the atrial cavity. The terminal 
sections of the paired oviducts and a section of the common oviduct 
are equipped with strongly eosinophilic shell glands. 

The copulatory bursa (6) is a large sac with a lobed outline. It 
connects posteriorly with a duct of almost uniformly wide diameter, 
the bursa stalk (bd). The duct runs first posteriorly, to the left of 
the penis bulb, then curves ventrally and opens, from the dorsal side, 
into the narrow terminal portion of the atrium, close to the genital 
pore. The bursa stalk has a strong coat of intermingled circular and 
longitudinal muscle fibers. There is no histological differentiation 
into anterior and posterior sections of the bursa stalk. 

Taxonomic position.—Polycelis borealis is the second species of the 
genus Polycelis to be found on the North American continent. The 
other species, P. coronata (Girard), reported from Wyoming and South 
Dakota (Hyman, 1931), resembles it closely in external appearance 
and probably cannot be distinguished from it in life. In both species, 
the shape of the anterior end is very similar. The auricles of P. 
borealis are perhaps a trifle longer and more pointed than those of P. 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 6 


Sketches of living specimens of the four Alaskan triclad species, all 10: 1. Phagocata 
nivea; 2. Polycelis borealis; 3. Dendrocoelopsis piriformis, striped form; 4. Dendrocoelop- 
sis alaskensis. 


FRESH-WATER TRICLADS OF ALASKA—KENK 171 


coronata. Hyman (1931, p. 126) states that in P. coronata the 
curved band of eyes narrows as it crosses the base of the auricles; 
no such narrowing has been seen in P. borealis. These differences 
are, however, insignificant and do not permit a clear separation of 
the two forms. The same may be said of other species of the genus, 
which likewise could be confused with the American forms in life: 
the Japanese species, P. auriculata and P. karafto; and P. schmidti 
occurring both in Kamchatka and in Japan. These forms differ 
from each other more clearly in their anatomical characters. 


| 
l 
| 
5 
Ficure 22.—Polycelis borealis, diagram of the copulatory organs in longitudinal section, 


X 70. a, atrium; b, bursa; bd, bursa stalk; de, ejaculatory duct; 0, mouth; ode, common 
oviduct; pg, genital pore; vd, vas deferens; vs, seminal vesicle. 


To gain a better insight into the systematic relations of Poly- 
celis borealis to other species of the same genus, it is necessary to review 
the present state of the systematics of the genus. The genus Poly- 
celis has a muscular pattern of the pharynx typical of the family 
Planariidae; the oviducts unite, without embracing the bursa stalk, 
to form an unpaired terminal oviduct; the testes are situated in the 
anterior part of the body only; the male atrium is not surrounded 
by radial muscle plates; and the eyes are numerous. The genus 
thus defined (Kenk, 1930) has today about twenty species. It may 
be subdivided into subgenera on the basis of structural characters 
of the copulatory apparatus. 

Subgenus Polycelis, lacking adenodactyls and lacking an excessive 
development of the muscle coat of the male atrium. This subgenus 
includes Sorocelides Sabussowa (1929, p. 521) and Polycelidia Zabu- 
sova (1936, p. 152), both described as distinct genera. The subgenus 
comprises the following species: 


238538—53——_2 


172 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Polycelis nigra (O. F. Miller), Europe P. polyopis Zabusova (1936), Kam- 


P. receptaculosa (Livanov and Zabusova, chatka 
1940), Teletskoe Lake in the Altai P. karafto Ijima and Kaburaki (1916), 
Mountains Sakhalin 
P. eburnea (Muth, 1912), Aral Sea P. sapporo (Ijima and Kaburaki, 1916), 
region Japan 
P. tibetica Hyman (1934), Tibet P. coronata (Girard, 1894), Wyoming 
P. koslowi (Zabusov, 1911), Tibet and South Dakota 
P. elongata (Sabussowa, 1929), Kam- 
chatka 


Subgenus Seidlia Zabusov (1911, suppl., p. 7), distinguished by an 
extraordinarily thick muscle zone surrounding the male atrium. 
Zabusov’s (1916, p. 273) genus Rjabuschinskya likewise belongs here. 
Species of the subgenus: 


Polycelis sabussowi (Seidl, 1911), in- P. schmidti (Zabusov, 1916), including 


cluding Seidl’s species Sorocelis P. ijimai Kaburaki (1922), Kam- 
sabussowi, S. lactea, S. stummeri chatka, Kurile Islands, and Japan 
(cf. Kenk, 1936), Turkestan P. auriculata Ijima and Kaburaki (1916), 
P. relicta (Sabussowa, 1929), Kamchatka Japan 
P. eurantron (Zabusova, 1936), Kam- 
chatka 


The third subgenus, Jjimia Bergendal (1890, p. 326), is charac- 
terized by possessing solid adenodactyls: 


Polycelis felina (Dalyell), including P. linkot Zabusov (1901), Onega Lake 


P. cornuta Johnson, Europe and in European Russia 

North Africa P. oculi-marginata (Palombi, 1931), New 
P. tenuis Ijima, Europe and western Guinea (see Beauchamp, 1947) 

Asia 


Two species of the genus Polycelis are too incompletely known to 
permit their assignment to either the subgenus Polycelis or Seidlia: 
P. eudendrocoeloides (Sabussowa, 1929) from the Kamchatka Penin- 
sula and P. tibetica (Zabusov, 1911, p. 349) from Tibet (P. tibetica 
Hyman, 1934, will have to be renamed if it should prove to be different 
from Zabusov’s species). 

Polycelis borealis is clearly a member of the subgenus Polycelis. 
It differs from P. nigra in having well-developed auricles; from P. 
elongata, P. polyopis, and P. sapporo, in the arrangement of the eyes; 
and from P. coronata and P. receptaculosa in the structure of the male 
copulatory organ. The reproductive system of P. karafto has not been 
described adequately and thus does not permit a comparison with 
that of the Alaskan species. The remaining species of the subgenus, 
P. eburnea, P. koslowi, and P. tibetica Hyman, two of which are known 
only from the study of preserved specimens, agree with P. borealis 
in being pigmented forms with prominent auricles and in having 
a similar anatomy of the copulatory organs. The four species are 


FRESH-WATER TRICLADS OF ALASKA—KENK 173 


undoubtedly very closely related; whether some of them are identical 
or are the same as the Alaskan form cannot be established on the basis 
of present knowledge. The Alaskan Polycelis, therefore, is described 
as a new species. 

Holotype.—On three slides, USNM 22333, clear spring on the road 
from Palmer to Willow, 20.1 miles from Palmer, altitude 3,800 feet, 
Aug. 9, 1950. 

Distribution and ecology.—Polycelis borealis is a very common species 
occurring in mountain streams in the southern part of Alaska (Alaska 
Range, Talkeetna Mountains, Chugach Range). Typically it 
inhabits clear, cold, fast-running waters. It has also been found, 
however, in several small mountain lakes which connect with streams 
in which the species lives. It has not been observed in silt-bearing 
glacier streams. 

The typical temperature range of the habitats was between 3.0° 
and about 15° C. (August 1950). Temperatures above 15° C. (up 
to 22.4° C.) were encountered only rarely and only in habitats which 
presumably have great diurnal temperature amplitudes (shallow lakes, 
small exposed streams). 

The great majority of the specimens collected was asexual. Sexually 
mature animals were seen in only 6 of the 31 localities in which the 
species was found.. In some localities, a large percentage of the speci- 
mens exhibited freshly regenerated heads or posterior ends, indicating 
that vivid asexual reproduction was taking place. 

Clear springs on the road from Palmer to Willow, 20.1 miles from Palmer, 
altitude 3,800 feet, Aug. 9, 1950, water temperature 3.0° C., numerous specimens, 
five mature (holotype). 

Clear, fast mountain stream crossing Mount McKinley Park road, 5.5 miles 
from the railroad station; Aug. 5, 1950. 

Streams on road from Palmer to Willow 11.9, 18.0, 19.2, 20.2, 21.6, 22.1, 22.8, 
and 34.0 miles from Palmer, and Ice Lake, 20.4 miles from Palmer, Aug. 9, 1950. 

Streams on Glenn Highway (Anchorage to junction with Richardson Highway 
near Glenallen) at mileposts 33.1, 33.9, 43.5, 64.8, 89.4, 98.8, and 117.1; Aug. 11 
and 13, 1950. 

Streams on road from Anchorage to Potter 2.9 (Campbell Creek), 10.3, and 10.6 
miles from the city limits of Anchorage, Aug. 12, 1950. 

Streams on Richardson Highway (section between Valdez and Glenallen) at 
mileposts 20.5, 25.5 (22.4° C.), 26.3 (see pl. 7), 29.0, 37.3, 54.7, 62.6 (see pl. 7), 


81.0 (Squirrel Creek), and 87.5 (Rock Creek), and lake at milepost 27.3 (22.1° C.), 
Aug. 15, 1950. 


Polycelis borealis was also identified in samples of triclads collected 
by Dr. Reece I. Sailer in July and September 1948. The following 


localities were represented: Richardson Highway, mileposts 8.3, 187.5, 
192.9, 209.7, 223.6 and 239.9, and Glenn Highway, milepost 117.1. 


174 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Family DENDROCOELIDAE 


Genus Dendrocoelepsis Kenk 


Dendrocoelopsis piriformis, new species 


Figure 23; Pate 6, Ficure 3 


Description.—Sexually mature specimens of this rather broad and 
plump species measure up to 15 mm. in length and 3 mm. in width. 
In quietly gliding animals, the head is truncated, with a convex 
frontal margin (grasping organ). There is a visible subterminal 
depression corresponding to the adhesive surface of the grasping 
organ or sucker. The rounded lateral corners of the head are formed 
by the auricles, which protrude only little laterally, thus causing an 
insignificant narrowing (neck) to appear behind them. Behind the 
head, the body margins widen gradually. The greatest width is 
reached at the level of the mouth. Behind that level, the body tapers 
again to a bluntly pointed posterior end. When the animal is at 
rest, the body appears short and wide, about pear-shaped, the body 
margins often forming a wavy or irregular contour. Young specimens 
are more slender and their lateral margins are more nearly parallel 
when they are in gliding motion. At rest, however, they assume the 
same pyriform shape as the adults. In short, the habit of the species 
may be described as resembling that of European and Asiatic repre- 
sentatives of the genus Bdellocephala Man, with which it has in com- 
mon the broad shape and the anterior grasping organ. 

The two eyes are situated on the dorsal side of the head. Their 
distance from each other amounts to somewhat more than one-third 
the width of the head at the level of the eyes. The distance of each 
eye from the lateral margin is smaller than the distance from the 
frontal margin. 

The color of the dorsal side is usually a cloudy brown or dark 
brownish gray. In some specimens, the pigment is arranged in 
definite longitudinal stripes: one sharply marked median stripe flanked 
on each side by a light (usually yellow) band; laterad to this band the 
body darkens, losing its pigment again near the lateral margin. In 
other specimens, lacking the two light dorsal bands, the back may be 
either uniformly pigmented or may show a more or less distinct darker 
band along the midline. Young specimens and striped adults clearly 
show the finer disposition of the pigment in small rounded dots, each 
dot apparently corresponding to an individual pigment cell. 

The pigment pattern of the dorsal side of the head is quite charac- 
teristic. A very dark field between the eyes extends anteriorly to 
both sides of the grasping organ. The area above the organ is unpig- 
mented and white, and so are the two elongated ocular areas antero- 


FRESH-WATER TRICLADS OF ALASKA—KENT 175 


laterad to the eyes. A pair of indistinct, converging streaks of lighter 
coloration may run posteriorly from the lateral angles of the head. 

The ventral side is light gray. 

The pharynx is situated, in mature animals, behind the middle of 
the body; the mouth, at the beginning of the last third of the body; 
and the genital aperture, midway between the mouth and the hind end. 

The normal locomotion of the animal is quiet gliding. When dis- 
turbed, it may attach itself to the substratum (apparently with the 
marginal adhesive zones) or move by “‘crawling.’”’ Young specimens 
do not crawl as readily as do adult ones. 

The grasping organ, or sucker, in living animals, appears as a well- 
marked bulge on the frontal margin of the head, showing a concave 
ventral depression. In preserved specimens, the frontal margin is 
generally curved ventrally and the site of the organ forms a thick, 
grooved rim. Anatomically, the organ consists of glandular and 
muscular elements. The subterminal adhesive surface is covered with 
an epithelium devoid of rhabdites and pierced by numerous gland 
ducts filled with a granular, eosinophilic secretion. The cell bodies 
of the glands are scattered through the mesenchyme of the anterior 
half of the prepharyngeal region, particularly above the intestine. 
The gland ducts run anteriorly in dense bundles and, in their terminal 
sections, are thickly swollen with secretion. The muscular system of 
the organ, which could not be analyzed in detail on account of the 
density of the glandular structures, has fibers attached to the adhesive 
surface, serving presumably as retractors. 

The rather short and thick pharynx is structurally typical of the 
family Dendrocoelidae; its internal muscular zone consists of inter- 
mingled circular and longitudinal fibers. 

Auricular sense organs are represented by two bands of sensory 
epithelium extending posteriorly from the sides of the frontal margin. 
The cells of this epithelium are less tall than are those of the sur- 
rounding body epithelum; they lack rhabdites and are approached by 
nerve fibers from the underlying mesenchyme. The location of these 
organs corresponds to the two light, longitudinal streaks on the sides 
of the head seen in the living animal. 

The testes are of moderate size, numerous, densely packed, and 
occupy the dorsal half of the mesenchyme, generally above the intes- 
tinal branches. They are arranged in two wide areas on both sides 
of the midline, extending from the level of the ovaries close to the 
posterior end. 

The ovaries, situated at the level of the second pair of branches of 
the anterior intestinal trunk, show no structural peculiarities. 

The genital pore (pg) leads immediately into two cavities, the male 
atrium (am) and, to the right and somewhat posteriorly, the duct of 


176 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


the copulatory bursa (bd). No common atrium is_ present, 
and both the papilla of the penis and the opening of the 
oviduct are situated in the “male” atrium. The atrium is a conical 
cavity, wide anteriorly and tapering toward the genital pore. It is 
lined with a cubical epithelium, below which occurs a thin layer of 
fine circular muscle fibers and a thicker layer of coarser longitudinal 
fibers. 

The penis consists of a spherical, muscular bulb and an elongated 
papilla. The penis bulb is differentiated into a wide peripheral 
muscular zone and a more central parenchymatic zone which contains 
a cavity, the seminal vesicle (vs). The vesicle is lined with a glandu- 
lar epithelium and its wall forms villuslike projections. The two 
vasa deferentia (vd) penetrate the penis bulb from the anterolateral 
sides and open into the seminal vesicle on two prominent conical 
papillae projecting from the anterior wall of the vesicle a short dis- 
tance from each other. 

The papilla of the penis is finger-shaped, tapering toward the tip, 
and is highly muscular. It is covered with a thin cubical epithelium. 
Below the epithelium there is, in the basal portion of the papilla, a 
thin layer of longitudinal muscles, below which lie a stronger circular 
layer and a second longitudinal layer. In the distal part of the 
papilla, the external longitudinal layer is lacking. The axis of the 
papilla is pierced by the ejaculatory duct (de) which leads from the 
seminal vesicle to the tip of the papilla. The duct has a cubical 
epithelium and a strong coat of longitudinal muscle fibers. 

The two oviducts approach the midline in the region of the copula- 
tory apparatus, the right one passing between the atrium and the 
bursa stalk, and unite behind and above the atrium. The common 
oviduct (odc) proceeds ventrally, then curves anteriorly, and empties 
into the terminal part of the atrium close to the genital pore. 

The copulatory bursa (b) is a lobed sac lined with a tall glandular 
epithelium. The distal parts of the epithelial cells are filled with 
fine eosinophilic granules. The duct or stalk of the bursa (bd) is dif- 
ferentiated into a narrow anterior section with a weak muscular coat, 
which connects with the bursa, and a posterior wider section which 
bends ventrally and opens at the genital pore. The cells of the 
anterior section resemble those of the bursa in having similar granular 
inclusions. Those of the posterior section lack the inclusions and are 
ciliated. The muscle coat of the duct consists of intermingled longi- 
tudinal and circular fibers. 

Taxonomic position—I have placed the species in the genus 
Dendrocoelopsis established originally for a European species, D. 
spinosipenis (Kenk). The original definition of the genus was based 
on the following characters: fibers of the inner muscle zone of the 


FRESH-WATER TRICLADS OF ALASKA—KENK 177 


pharynx intermingled; no adenodactyl; penis papilla developed, penis 
bulb of simple structure; oviducts unite without embracing bursa 
stalk; zone of testes extending behind the level of the copulatory 
organs; anterior end with subterminal true sucker; eyes not numerous 
(Kenk, 1930). Subsequently Beauchamp (1932, p. 254) founded a 
new genus, Amyadenium, with two species, A. vandelt Beauchamp and 
A. brementi (Beauchamp), both from the Pyrenees. Two more 
species were reported by the same author in later papers, A. chattona 
Beauchamp (1949, p. 60), again from the Pyrenees, and A. garmieri 
Beauchamp (1950, p. 65), from central France. Beauchamp recog- 
nized the close relation of the new genus to Dendrocoelopsis, but 


VS 
: BNW Ze 


4 
| 


— oa 


o bl ouvd 


Ficure 23.—Dendrocoelopsis piriformis, diagram of the copulatory organs in longitudinal 
section, X 60. am, maleatrium; d, bursa; bd, bursa stalk; de, ejaculatory duct; 0, mouth; 
odc, common oviduct; pg, genital pore; od, vas deferens; vs, seminal vesicle. 


separated it from the latter on account of the absence of a highly 
complex grasping organ, or true sucker (1. e., an adhesive organ sepa- 
rated from the surrounding mesenchyme by a muscle layer). Hyman 
(1935) described a dendrocoelid species from Montana under the 
name of D. vaginatus. Again the main characters of the species 
coincide with those of D. spinosipenis with the exception of the 
grasping organ, which is of a simpler type. The new Alaskan species, 
D. piriformis, also falls clearly in the vicinity of the species enumer- 
ated, as does another species, D. alaskensis, the description of which 
follows on p. 178. Within this group of species there is a gradual 
differentiation of the grasping organ, which is absent in D. alaskensis, 
present as a moderately developed adhesive organ in D. vaginata, 
D. piriformis, A. brementi, A. vandeli, A. chattoni, and A. garmieri, 
and as a more highly developed and muscular sucker in D. spinosipenis. 
A similar wide variation of the structure of the anterior grasping organ 


178 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


is seen in the genus Dendrocoelum Orsted where, however, true suckers 
are not known. In general, the taxonomic value of adhesive organs 
in triclads appears to be subordinated to that of other anatomical 
structures. I therefore tentatively modify the definition of the 
genus Dendrocoelopsis by omitting the presence of a sucker as a 
generic character, to include the species described as Dendrocoelopsis 
and Amyadenium. 

Dendrocoelopsis pirvformis differs from the other members of the 
genus (in its wider extent) in several characters: It is pigmented, 
whereas the others lack pigment and appear white; the presence of 
two eyes differentiates it from the three blind species, D. vandeli, D. 
brementi, and D. garmieri and from the many-eyed D. chattoni; and 
the dorsal position of the testes separates it from D. spinosipenis, D. 
vaginata, D. garmeri, and D. alaskensis. Apart from these most con- 
spicuous characters, the structure of the male copulatory organ of D. 
piriformis is distinctive. 

Holotype.—On five slides, USNM 22334, Moose Creek, on Glenn 
Highway, milepost 186, near Alaska Road Commission camp, Aug. 
14, 1950. 

Distribution and Ecology.—Dendrocoelopsis piriformis is a eury- 
thermic species and inhabits lakes and their outlets in the southern 
part of Alaska and also occurs in a lake near Umiat. 

Long Lake, on Glenn Highway, milepost 85.9, Aug. 11, 1950, clear water, 
17.8° C. (near bank) ; under stones, several specimens, two of them mature. 

Lake on Glenn Highway, milepost 88.1, Aug. 11, 1950, water temperature 
varying with depth (near bank, 19° C.); under stones, several specimens, one 
mature. 

Lake on Glenn Highway, milepost 23.5 (see pl. 8), Aug. 13, 1950, clear water, 
23.6° C. (near bank); several immature specimens. 

Stream crossing Glenn Highway at milepost 147.2, outlet of Snowshoe Lake, 
Aug. 14, 1950, moderate current, water somewhat colored, 17.0° C.; one young 
specimen, under a stone. 

Moose Creek, on Glenn Highway, milepost 186, near Alaska Road Commission 
camp, Aug. 14, 1950, fast, clear stream, 16.2° C.; under stones, two mature speci- 
mens (holotype). 

Pippin Lake, on Richardson Highway, milepost 84.4, Aug. 15, 1950, shallow 
water near bank, 20° C.; under stones, many specimens, two of them mature. 

Fresh-water lake near Umiat, collected by P. F. Scholander, Aug. 15, 1948; 


18 specimens, majority belonging to the plain form, 1 with distinet stripes, 5 
sexually mature (USNM 23678). 


Dendrocoelopsis alaskensis, new species 
Ficure 24; Puate 6, FIGuRE 4 


Description.—I had only limited material of this species and none 
of the animals was fully mature. The largest specimen measured 20 
mm. in length and 4 mm. in width. 

The anterior end is slightly lobed, with a convex frontal margin 


Eee 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 7 


ee ont oe : ai “ tie, owe a a a 
cote a — er ES Mee Lanthanide Satna . a 


Top: Beaver pond at milepost 62.6 of Richardson Highway, Alaska. Below the beaver 
dam, Polycelis borealis under stones. Bottom: Stream flowing along Richardson High- 
way, Alaska, at milepost 26.3. On the undersides of stones, Polycelis borealis. 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 8 


Top: Stream crossing Steese Highway, Alaska, at milepost 84.0. Habitat of Phagocata 
nivea and Dendrocoelopsis alaskensis. Bottom: Lake at milepost 23.5 of Glenn High- 
way, Alaska. Under stones near bank, Dendrocoelopsis piriformis. 


FRESH-WATER TRICLADS OF ALASKA—KENK 179 


and a pair of rounded auricles protruding both anteriorly and later- 
ally. No distinct adhesive or grasping organ is developed. There is 
a gentle narrowing of the head (neck) behind the auricles. Behind 
the head, the width of the body gradually increases until the greatest 
width is reached. The posterior end is bluntly pointed. 

The body lacks pigment, being white except for the contents of 
the intestine, which may show through the body wall. 

There are two principal eyes. The distance between them amounts 
to about one-third the width of the head at the level of the eyes. 
The distance of each eye from the frontal margin is equal to, or 
slightly larger than, the distance from the lateral margin. Additional, 
supernumerary, eyes were seen in a few individuals at short distances 
either anterior or posterior to the principal eyes. 

The species bears a striking resemblance to a European species, 
Dendrocoelum nausicaae Schmidt of the Balkan Peninsula. It is 
generally associated with another triclad inhabiting the same streams 
in Alaska, Phagocata nivea, from which it may be distinguished by 
its larger size and by the position of the eyes. 

A distinct adhesive organ is not seen in living animals. In histologi- 
cal sections, a transverse band, pierced by numerous openings of 
eosinophilic glands, is found below the frontal margin of the head. 
The nature of the glands and the local differentiation of the epithelium 
correspond entirely to the structure of the submarginal adhesive zone 
which is seen in this species, as well as in other triclads, bordering the 
lateral margins of the body. The frontal adhesive area may, therefore, 
be interpreted as an extension of the lateral adhesive zone. It is 
somewhat wider than the lateral one and has no muscular differen- 
tiations such as are typical of true grasping organs. The continuity 
of the two zones is interrupted by a short gap on each side of the 
head below the auricle. 

The internal muscle zone of the pharynx consists of a layer of inter- 
mingled circular and longitudinal fibers. This character places the 
species in the family Dendrocoelidae. The anterior intestinal trunk 
bears a fairly large number, 21 to 24 pairs, of lateral branches, or 
diverticula. 

Of a total of eight individuals of this species at my disposal, five 
were young, two showed primordia of genital structures, and only one 
had the principal parts of the reproductive system developed, though 
not fully differentiated histologically. The description of the genital 
organs, therefore, must be given with certain reservations. 

The testes are predominantly ventral, situated mainly below the 
level of the intestinal diverticula, occasionally extending farther 
dorsally between the diverticula. No clearly recognizable testes were 
seen behind the level of the mouth; they may, however, appear there 


180 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 103 


when the animal matures completely. The ovaries are situated 
behind the third or fourth lateral branch of the anterior intestinal 
trunk. 

The genital pore (pg) leads into an undivided cavity, the genital 
atrium (a), which is narrow at the pore and expands anteriorly. The 
narrow posterior part receives the outlet of the copulatory bursa and, 
anteriorly to it, the opening of the common oviduct. 

The penis has a fairly large, spherical bulb and a plump and short 
papilla. The bulb contains a cavity with irregular outline, the seminal 
vesicle (vs), into which the two vasa deferentia (vd) open separately. 
The cavity of the bulb continues into the broad papilla and opens at 
its tip. The bulb has the usual coat of muscle fibers arranged in 
concentric layers. The papilla, which projects only little into the 
genital atrium, has two muscle layers underlying the outer epithelium, 
a circular layer and a longitudinal one. No glandular structures are 
differentiated in my specimen. 

The two oviducts unite, without embracing the bursa duct, dorsally 
to the genital atrium. The common oviduct (odc) opens into the 
posterior, narrow part of the atrium from the dorsal side. 

The bursa (b) is, in my specimen, a rather small sac with a narrow 
lumen. The bursa duct (6d) runs posteriorly, above the penis, to a 
level behind the genital pore. There it turns abruptly toward the 
ventral side. Its terminal portion is considerably wider than the 
anterior part of the duct and opens from the posterodorsal side into 
the genital atrium close to the genital aperture. In full maturity, the 
widened section of the bursa duct probably represents a histologically 
distinct vagina. 

Taxonomic position.—The systematic relations of Dendrocoelopsis 
alaskensis are discussed together with those of the preceding species, 
D. piriformis. D. alaskensis is distinguished from all other species 
of the genus by the lack of a grasping organ. 

Holotype. On seven slides, USNM 22335, creek crossing Elliot 
Highway at milepost 31.0, July 24, 1950. 

Distribution and ecology.— Dendrocoelopsis alaskensis is an inhabitant 
of cool, fast streams of the White Mountains and usually shares its 
habitat with Phagocata nivea. It is a stenothermic and rheophilic 
species. 

Clear spring and creek on Steese Highway, milepost 82.5, at the Alaska Road 
Commission camp, July 21, 1950; one immature specimen, under a stone. 

Stream crossing Steese Highway at milepost 84.0 (see pl. 8), altitude 2,700 
feet, July 21, 1950; two immature specimens, under stones, near liver bait. 

Creek crossing Elliot Highway at milepost 31.0, July 24, 1950, water tempera- 
ture 3.2° C.; under stones, five specimens, two of them with sexual structures 
(holotype). 


FRESH-WATER TRICLADS OF ALASKA—KENK 181 


ae . 
yoy 
SY 
l 


| 

| 

| 
tps, 
ph b ed vs a odcpg bd 
Ficure 24.—Dendrocoelopsis alaskensis, diagram of the copulatory organs in longitudinal 
section, X 70. a, genital atrium; b, bursa; bd, bursa stalk; odc, common oviduct; 


pg, genital pore; ph, pharynx (extended through the mouth); od, vas deferens; 
os, seminal vesicle. 


Zoogeographic conclusions 


The occurrence in Alaska of four endemic species of triclads poses 
several interesting questions. It is known that fresh-water triclads 
are most abundantly represented in the northern Temperate Zone 
and that the number of species declines toward both the Arctic and 
the Tropical Zones. The paucity of the triclad fauna at high lati- 
tudes has frequently been attributed to the effects of glaciation in 
rather recent geologic time. During the glacial period, when huge 
ice masses covered great parts of the northern hemisphere, all fresh- 
water life over wide areas must have vanished. After the glaciers 
had receded, the areas were gradually repopulated by species entering 
them from adjacent territories. The pattern of distribution of 
freshwater triclads in Europe shows evidence of a definite succession 
of species which migrated into the previously glaciated areas in the 
postglacial period (cf. Thienemann’s 1950 summary of the pertinent 
literature). 

It is well known, however, that the greater part of Alaska was 
not covered with ice in the glacial period. Geologic evidence of 
glaciation has been found only in the Brooks Range, the Alaska 
Range (and areas south of it), and in small isolated spots in the 
Yukon and Kuskokwim River Valleys while the remaining surface 
of Alaska remained free of ice (cf. Geological Society of America, 


182 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 103 


1945). We are thus justified in assuming that the fresh-water life 
of Alaska was not completely destroyed even at the peaks of glacia- 
tion when practically all Canada and a considerable part of the 
United States were covered with ice caps. 

The Alaskan fresh-water triclads show no close relationships with 
the present North American triclad fauna inhabiting the midwestern 
plains and the eastern and southern areas of the continent. A 
comparison with the fauna of the Rocky Mountains is not possible 
at present, since the West of the United States and of Canada is 
almost unexplored with regard to triclads and to lower aquatic in- 
vertebrates in general. In any event it appears highly improbable 
that Alaska was populated by species migrating to it from the south 
or east. 

On the other hand, the relations of the Alaska triclads to the 
fauna of Eurasia are unmistakable. Though, according to our 
present knowledge, none of the Alaskan triclads is specifically identi- 
cal with any Asiatic form, two species, Phagocata nivea and Polycelis 
borealis, are very closely related to Asiatic species of the same genera. 

Figure 25 shows the geographic range of the genus Polycelis as it 
is known at present. The individual dots on the map represent 
either single records or groups of neighboring localities where species 
of the genus have been found. In interpreting the map it is to be 
kept in mind that not all geographical areas are equally well investi- 
gated with regard to the occurrence of lower invertebrates. Europe, 
Japan, and the eastern parts of the United States are comparatively 
well known while our knowledge of the Asiatic, Western American, 
and African triclads is still rather deficient. A study of the map 
suggests that Polycelis is primarily a Eurasian genus. It appears 
to have extended its range, in some earlier geological period, to the 
northern rim of Africa. In a similar way it may have migrated into 
Alaska at the time of the cenozoic land bridges and may have pene- 
trated south along the Rocky Mountains. It is highly probable 
that Polycelis will be found, in the future, more widely distributed 
in the Rockies than present collection records indicate. 

The repeated emergence of a land bridge between Alaska and the 
eastern tip of Siberia is generally accepted by geologists and paleon- 
tologists. There is ample evidence of an exchange of faunal elements 
between the two continents (cf. Simpson, 1940 and 1947). The most 
recent corridor must have existed during the glacial stages of the 
Pleistocene. The volume of ice masses accumulated over vast areas 
of the northern hemisphere has been estimated conservatively at 
34 to 42 million cubic kilometers. The corresponding depletion of 


‘4 The presence of Polycelis oculi-marginata (Palombi) in New Guinea is a zoogeographic enigma (cf. Beau- 
champ, 1947). 


183 


FRESH-WATER TRICLADS OF ALASKA—KENK 


"syaIKjog Snuas ay} JO UOINISIp spIMp]IoM—'¢Z AWNOL] 


184 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 103 


the ocean must have lowered the sea level 50 to 90 meters below the 
present level (Daly, 1934, pp. 41-50). The depth of the eastern part 
of the Bering Sea (within the perimeter passing through the eastern 
projection of Siberia, the Seward Peninsula of Alaska, and Nunivak 
and St. Lawrence Islands) averages less than 40 meters. Thus the 
ocean bottom between Alaska and Asia must have emerged with each 
glacial stage, forming a wide connection between the two continents. 
This process may have been enhanced by the uplifting of the area 
along the margin of the ice cap, brought about by plastic or elastic 
deformation of the earth crust under the weight of the ice masses. 

A land bridge between Alaska and Asia must have persisted for 
prolonged periods of time. Simultaneously, Alaska was discon- 
nected from other inhabitable areas of North America by broad ice 
fields. 

In view of the geological history of Alaska we may conclude that 
the present Alaskan fresh-water triclads are the remnants or successors 
of triclads that lived in Alaska in the glacial times and perhaps even 
in preglacial periods, and that they are, in all probability, of Asiatic 
origin. It may further be assumed that some Alaskan triclads have 
extended their range during the postglacial period and proceeded 
southeast along the Rocky Mountains.’ The occurrence of a species 
of Polycelis in Wyoming and South Dakota may have resulted from 
this migration. More light would be thrown on this question by a 
more thorough study of the triclad fauna of the Canadian and 
American Rockies. 


Literature cited 
ARNDT, WALTHER 
1922. Untersuchungen an Bachtricladen. Ein Beitrag zur Kenntnis der 
Paludicolen Korsikas, Rumianiens und Sibiriens. Zeitschr. fiir wiss. 
Zool., vol. 120, pp. 98-146, pl. 4. 
Bazigatova, A. fA. 
1947. Turbellaria-Triclada vostochno! Sibiri i Pribafkal’fa. Doklady Akad. 
Nauk SSSR, new ser., vol. 55, pp. 671-672. 
BEAUCHAMP, PAUL DE 
1932. Biospeologica. LVI. Turbellariés, Hirudinées, Branchiobdellidés. 
Deuxiéme série. Arch. Zool. Expér, et Gén., vol. 73, pp. 113-380, 
pls. 6-8. 
1939. Lasystématique et l’éthologie des Fonticola (Turb. Triclades). Vé&stnik 
Ceskoslov. Zool. Spoleénosti v Praze, vol. 6-7, pp. 91-96. 
1947. Observations sur quelques Turbellariés du Musée royal d’Histoire 
naturelle de Belgique. Bull. Mus. Roy. Hist. Nat. Belgique, vol. 
23, No. 33, 11 pp. 
1949. Biospeologica. LXIX. Turbellariés (troisitme série). Arch. Zool. 
Expér. et Gén., vol. 86, Notes et Revue, pp. 50-65. 
1950. Nouvelles diagnoses de triclades obscuricoles. Bull. Soc. Zool. France, 
vol. 75, pp. 65-70. 


5I wish here to correct my assumption in a previous paper (Kenk, 1943, p. 6) that all Canadian triclads 
had entered Canada from the south. 


FRESH-WATER TRICLADS OF ALASKA—KENK 185 


BERGENDAL, D. 
1890. Studien tiber nordische Turbellarien und Nemertinen. Ofv. Vet.- 
Akad. Forh., 1890, pp. 323-328. 
Daty, REGINALD ALDWORTH 
1934. The changing world of the ice age. xx1t+271 pp., 6 pls. New Haven. 
GEOLOGICAL SocrETY OF AMERICA 
1945. Glacial map of North America. Special Papers, No. 60. 
GIRARD, CHARLES 

1894. Recherches sur les Planariés et les Némertines de 1’Amérique du Nord. 

Ann. Sci. Nat., sér. 7 (Zool.), vol. 15, pp. 145-310, pls. 3-6. 
Hyman, LisBiE H. 

1931. Studies on the morphology, taxonomy, and distribution of North 
American triclad Turbellaria. III. On Polycelis coronata (Girard). 
Trans. Amer. Micr. Soc., vol. 50, pp. 124-135. 

1934. Report on triclad Turbellaria from Indian Tibet. Mem. Connecticut 
Acad. Arts Sci., vol. 10, No. 2, pp. 5-12, pls. 1-2. 

1935. Studies on the morphology, taxonomy, and distribution of North 
American triclad Turbellaria. VI. A new dendrocoelid from 
Montana, Dendrocoelopsis vaginatus n. sp. Trans. Amer. Micr. 
Soc., vol. 54, pp. 338-345. 

1937. Studies on the morphology, taxonomy, and distribution of North 
American triclad Turbellaria. VII. The two species confused 
under the name Phagocata gracilis, the validity of the generic name 
Phagocata Leidy 1847, and its priority over Fonticola Kom4rek 1926. 
Trans. Amer. Micr. Soc., vol. 56, pp. 298-310. 

Isma, I., and Kasurakxr, T. 

1916. Preliminary descriptions of some Japanese triclads. Annot. Zool. 

Japon., vol. 9, pp. 153-171. 
KaBuRAKI, TOKIO 

1922. On some Japanese freshwater triclads; with a note on the parallelism 
in their distribution in Europe and Japan. Journ. Coll. Sci., Imp. 
Univ. Tokyo, vol. 44, No. 2, 71 pp., 1 pl. 

KeENK, RoMAN 

1930. Beitrage zum System der Probursalier (Tricladida paludicola), III. 
Zool. Anz., vol. 89, pp. 289-302. 

1936. Bemerkung zur Trikladenfauna Turkestans. Zool. Anz., vol. 115, 
pp. 76-80. 

1943. Notes on the planarian fauna of Canada. Canadian Field-Nat., 
vol 57, pp. 5-6. 

Livanov, N. A., and Zasusova, Z. I. 

1940. Planarii basseina Telefskogo ozera i novye dannye o nekotorykh 
drugikh sibirskikh vidakh (Paludicoles du Lac Teletzkoé et nouvelles 
données sur quelques formes sibériennes). Trudy Obshch. Estes- 
tvoisp. pri Kazan. Univ., vol. 56, pp. 83-159. 

Mor, ANTON 

1912. Beitrige zur Kenntnis der Gattung Sorocelis Grube. Mitt. Naturw. 

Ver. Steiermark, vol. 48, pp. 381-410. 
PaLomBI, A. 

1931. Turbellari della Nuova Guinea. In: Résultats scientifiques du 
voyage aux Indes Orientales Néerlandaises de LL. AA. RR. le 
Prince et la Princesse Léopold de Belgique, vol. 2, No. 8, 14 pp., 
1 pl. (Mém. Mus. Roy. Hist. Nat. Belgique, hors série). 


186 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Sasussowa, Z. (see also ZaBusova, Z. I.) 

1929. Die Turbellarien der Kamtschatka-Halbinsel nach den Sammlungen 
der Rjabuschinsky-Expedition 1908-1909. Zool. Jahrb., Abt. 
Syst., vol. 57, pp. 497-536, pl. 4. 

SeripL, Hernricn H. 

1911. Beitrage zur Kenntnis centralasiatischer Tricladen. Zeitschr. fiir 

wiss. Zool., vol. 98, pp. 31-67, pls. 5-7. 
Simpson, GEORGE GAYLORD 

1940. Mammals and land bridges. Journ. Washington Acad. Sci., vol. 
30, pp. 137-163. 

1947. Holarctic mammalian faunas and continental relationships during 
the Cenozoic. Bull. Geol. Soc. Amer., vol 58, pp. 613-687. 

THIENEMANN, AUGUST 

1950. Verbreitungsgeschichte der Siisswassertierwelt Europas; Versuch 
einer historischen Tiergeographie der européischen Binnengewaisser. 
XVI-+809 pp. Stuttgart. 

ZaBusov’”, I. P. (Sasussow, H.; ZasBoussorr, H.) 

1901. Zamsétki po morfologii i sistematikié Triclada. II. O planarifakh” 
Onezhskago ozera (Tricladenstudien. II. Zur Kenntniss der 
Tricladen des Onegasees). Protok. Zasféd. Obshch. Estestvoisp. 
Imp. Kazan. Univ., Prilozhenie No. 191, 18 pp. 

1911. Izslié@dovanifa po morfologii i sistematikfé planarif ozera Batkala. 
I. Rod” Sorocelis Grube (Untersuchungen tiber die Morphologie 
und Systematik der Planarien aus dem Baikalsee. I. Die Gattung 
Sorocelis Grube). Trudy Obshch. Estestvoisp. Imp. Kazan. Univ., 
vol. 43, No. 4, 422+8-+2 pp., 11 pls. 

1916. Rjabuschinskya schmidti n. g. n. sp., novy! vid” i rod” Tricladida 
paludicola iz’? Kamchatki (Rjabuschinskya schmidti n. g. n. sp., 
espéce et genre nouveau des Tricladida paludicola du Kamtchatka). 
Russ. Zool. Zhur., vol. 1916, pp. 273-286. 

Zapusova, Z. I. (Sasoussorr, Z.; see also SaBussowa, Z.) 

1936. Planarii Kamchatki (Les planaires paludicoles du Kamtchatka). 
Kazan. Gosud. Univ., Uchenye Zapiski, Zool., vol. 96, No. 7, pp. 
141-174, 


U. S. GOVERNMENT PRINTING OFFICE: 1983 


PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1953 No. 3323 


REVIEW OF THE INDO-PACIFIC ANEMONE FISHES, GENUS 
AMPHIPRION, WITH DESCRIPTIONS OF TWO NEW 
SPECIES. 


By Lreonarp P, ScHuttz 


During the summer of 1950 Dr. Arthur D. Welander, School o 
Fisheries, University of Washington, and I were engaged in studying 
reef fishes brought back from the Marshall Islands by the staff of the 
Applied Fisheries Laboratory of the University of Washington. 
Among this material was a specimen of anemone fish that we could 
not identify with any known species. After I returned to the U. S. 
National Museum, I reviewed the descriptions of all known species, 
compared that specimen with the numerous lots of Amphiprion in 
the National Museum, and found that it represented a new species. 
During March 1951, I studied the anemone fishes in the Museum 
of Comparative Zoology, Harvard University, and found another 
undescribed species from Mauritius. 

Descriptions and analyses of species referable to the genus Amphi- 
prion have been based on so few specimens, usually only one or two, 
that the problem of variability or constancy of the color pattern has 
been neglected. For most of the few hundred species among more 
than fifty fish families that I have studied in detail the basic color 
pattern has been observed to be fairly constant. It is of great value 
for the recognition of species, especially in the genus Amphiprion. 
Weber and de Beaufort (The fishes of the Indo-Australian Archipelago, 
vol. 8, pp. 330-348, 1940) recognized eight species, whereas we have 
distinguished fourteen and there may be others recognizable when 
larger series are compared and additional characters studied. Fin 
ray counts were made on various species and these data are recorded 
in table 1. 

Except for original descriptions, no attempt was made to include 
all references to species referable to the genus Amphiprion. Whenever 


238537—_53 187 


188 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


figures of species were found these have been included in the synonymy. 
Most descriptions and records are not in sufficient detail to assign 
them to the correct species without examining the material on which 
the records were based. 


Genus Amphiprion Bloch and Schneider 


Amphiprion Bloch and Schneider, Systema ichthyologiae ..., p. 200, 1801 
(genotype, Lutjanus ephippium Bloch). 

Prochilus (on Klein, 1775) Bleeker, Nat. Verh. Hollandsche Maatsch. Wet. 
Haarlem, ser. 3, vol. 2, No. 6, p. 20, 1877 (genotype, Lutjanus ephippium 
Bloch). 

Actinicola Fowler, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 12, p. 533, 
1904 (genotype, Lutjanus percula Lacepéde). 

Phalerebus Whitley, Mem. Queensland Mus., vol. 9, pt. 3, p. 216, 1929 (genotype, 
Prochilus akallopisos Bleeker). 


Key to the species of Amphiprion 


la. A white band (sometimes indistinct) along middorsal line from snout to 
dorsal origin or beyond along base of dorsal fin; total pectoral rays 17 to 
19; next to last dorsal spine about 1.3 to 1.5 in longest dorsal spine; no 
notable emargination in dorsal fin; scales on dorsal surface of head extend 

forward to a line between front of orbits. 
2a. No vertical pale bars; dorsal rays about X,18 or 19; anal about II,12 or 
13 spectoral MifstorlO.s ss, 2 eee ee ee A. akallopisos Bleeker 
2b. A single vertical pale bar about 2 scales wide from nape to subopercle; 
dorsal rays about X,16; anal about II,12 or 13; pectoral 17. 
A. perideraion Bleeker 

1b. No white band along middorsal line. 
3a. Caudal fin with pale or dusky roundish center posteriorly edged with 
black; outer edges of caudal fin white; second pale or white bar from 
rear of spiny dorsal fin to anus with a triangular anterior projection 
under depressed pectoral fin; tips of pelvics black; a broad white 
color bar on head and on caudal peduncle always present. Dorsal 
fin deeply indented at rear of spiny part, next to last dorsal spine 
contained three times in longest dorsal spine; scales on dorsal surface 
of head do not extend forward of nape..__-_-- A. percula (Lacepéde) 
3b. Color not as in A. percula. 

4a. Central part of caudal fin black; outer edges of caudal fin broadly or 
narrowly edged with white; second pale bar, if present, without 
any projection anteriorly; pale bar on head present; next to last 
dorsal spine contained 1.2 to 2.0 times in longest dorsal spine; no 

notable emargination in dorsal fin. 
5a. Second pale bar represented dorsally on body by an ovate white 
area that continues anterodistally on soft dorsal fin; this white 
area does not extend below midlengthwise axis of body and 
never to anus; first pale bar 10 to 12 scales wide; anal fin black, 
except distally edged with white; spiny dorsal black; pelvics 
black; pectoral pale, except basally blackish; next to last dorsal 
spine contained about 1.2 to 1.5 in longest dorsal spine; no 
notable emargination in dorsal fin; scales on dorsal surface of 

head extend forward to a line between rear of orbits. 

A. laticlavius Cuvier and Valenciennes 


INDO-PACIFIC FISH GENUS AMPHIPRION—SCHULTZ 189 


5b. Second pale bar continuous from dorsal part of body to region 
of anus. 
6a. Second pale bar broad, about 7 to 14 scales wide at level of 
lateral line (its width there contained 2 or less times in width 
of third black bar at level of lateral line) from whence it con- 
tinues posterodorsally on soft dorsal fin; pelvics dusky to 
blackish; spiny dorsal blackish; anal blackish, at least basally; 
scales on dorsal surface of head extend forward to a line 
between rear of orbits. 
7a. Caudal peduncle with broad white bar; pectoral fin pale dis- 
tally, basally dusky. 
A. chrysogaster Cuvier and Valenciennes 
7b. Caudal peduncle black; no white bar; black coloration of 
posterior part of body continues on central part of caudal 
fin; pectoral fin dusky in basal third_A. poiymnus (Linnaeus) 
6b. Second pale bar narrow, about 2 to 6 scales wide at level of 
lateral line, its width there contained 3.5 or more times in 
width of third black bar at level of lateral line; caudal fin black, 
narrowly edged with white; anal fin pale to blackish; scales 
on dorsal surface of head extend forward to over rear half of 
pupil; pelvics and pectorals pale; soft dorsal black, edged with 
white. 
8a. Second and third (peduncular) pale bars about 2 or 3 scales 
wide, their width at level of lateral line contained 7 to 
10 times in width of third black bar at level of lateral 
line; second pale bar not extending to distal edge of 
dorsal fin, ending on basal half of last dorsal spine; 
posterodorsal part of body blackish; anterior and ventral 
parts of body pale; spiny dorsal light dusky. 
A. tricinetus Schultz and Welander, new species 
8b. Second and third (peduncular) pale bars 4 to 6 scales wide, 
their widths at level of lateral line contained from 4 to 
6.5 times in width of third black bar at level of lateral 
line; second pale bar extends into distal half of dorsal but 
not quite to the edge of that fin; dorsal and posterior half 
of body blackish; anteroventral part of body pale; spiny 
dorsal dusky to blackish. 
A. mauritiensis Schultz, new species 
4a. Caudal fin plain pale or plain dusky, no black central blotch edged 
with white posteriorly; first pale bar on head present. 
9a. Second pale bar broad, about 7 to 10 scales wide at level of lateral 
line, and continuing to distal edge of spiny dorsal fin, thence 
posteriorly along distal edge of soft dorsal; caudal fin pale; 
caudal peduncle black, without the third pale bar, at least on 
adults; spiny dorsal, anal, and pelvic fins black; pectoral fin pale; 
next to last dorsal spine contained about 1.5 in longest dorsal 
spine; scales on dorsal surface of head extend forward to a line 
between. rearioh Oroite 2 on ese A. sebae Bleeker 
9b. Second pale bar, if present, not continuing to distal edge of dorsal 
fin; next to last dorsal spine contained 0.8 to 1.2 in longest dorsal 
spine; scales on dorsal surface of head extend forward to lines 
between rear edge of orbits to center of pupil. 
10a. First two pale bars typically present on adults. 


190 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


lla. First two pale bars broad, each about 5 to 7 scales wide at 
level of lateral line; third (peduncular) pale bar represented 
by a white bar, posteriorly edged with darkish, or the dark 
body color on anterior part of caudal peduncle may end 
abruptly, the white continuing on caudal fin; spiny dorsal 
black, dusky, or pale; soft dorsal and anal fins pale to black; 
pelvics pale to black, sometimes edged with black anteriorly; 
pectoral pale; next to last dorsal spine about 1.2 in longest 
dorsal spine_..__.__._A. xanthurus Cuvier and Valenciennes 
11b. First two pale bars narrow, second usually narrower than first, 
the latter 3 to 4.5 scales wide and the first 3 to 6 scales wide, 
at level of lateral line; caudal peduncle blackish, without 
third pale bar on adults, sometimes pale bar is present on 
young; dark color of caudal peduncle gradually fading into 
pale color of caudal fin; spiny dorsal pale to dusky, soft 
dorsal pale; anal fin pale; pectoral pale; pelvics pale, except 
anterior edge black; second from last dorsal spine contained 
from 1.0 to 1.2 times in longest dorsal spine. 
A. bicinctus Riippell 
10b. First pale bar usually present on head, about 4 to 6 scales wide at 
level of lateral line; second pale bar lacking, except sometimes 
on small young specimens. 
12a. Anal fin pale or partly pale; pelvics with some pale area. 
13a. Anal fin pale, except distally edged with fine black line; 
pelvics pale, except anterior edge black; breast pale; 
dorsal spines X, soft dorsal rays usually 17. 
A. ephippium (Bloch) 
13b. Anal fin pale or distally pale with basal half (or less) 
dusky to blackish; pelvics pale distally and blackish 
ventrally, or inner rays pale and outer rays broadly 
blackish, except distal tips, which are pale; dorsal spines 
IX or X (more often IX than X), soft dorsal rays usually 
POP Gees se ae a. ee ee A. frenatus Brevoort 
12b. Anal and pelvic fins black, no pale areas anywhere; dorsal 
spines X, soft dorsal rays usually 17. 
A. melanopus Bleeker 


Amphiprion akallopisos Bleeker 


Puate 9, Figure A 


Amphiprion akallopisos Bleeker, Nat. Tijdschr. Nederl.-Indié, vol. 4, p. 281, 
1853. 

Prochilus akallopisus Bleeker, Atlas ichthyologique .. . , vol. 9, pl. 400, fig. 3, 
1878. 

Phalerebus Whitley (new genus), Mem. Queensland Mus., vol. 9, pt. 3, p. 216, 
1929 (genotype, Prochilus akallopisus Bleeker=A. akallopisos Bleeker). 

This species is best recognized by the presence of a wide white 

band from snout along middorsal line of head, thence posteriorly 

along each side of base of dorsal fin, and ending on dorsal side of caudal 

peduncle; no white bars are present. Three specimens were studied: 

USNM 147130, from the Philippine Islands; USNM 82781, from the 

Fiji Islands; and MCZ 3308, from Sabang Bay. 


INDO-PACIFIC FISH GENUS AMPHIPRION—SCHULTZ 191 


Amphiprion perideraion Bleeker 
Puate 9, Figure B 


Amphiprion perideraion Bleeker, Nat. Tijdschr. Nederl-Indié, vol. 9, p. 437, 
1855.—Montalban, Bur. Sci. Manila Monogr. 24, p. 16, pl. 4, fig. 1, 1928 
(Philippine Islands). 

Prochilus perideraion Bleeker, Atlas ichthyologique ... , vol. 9, pl. 400, fig. 1, 


1878. 
?Amphiprion rosenbergi Bleeker, Acta Soc. Sci. Indo-Neerl., vol. 6, p. 16, 1859; 
Atlas ichthyologique . . . , vol. 9, pl. 402, fig. 2, 1878. 


This species is best recognized by the narrow white band on the 
middorsal line of head, beginning between front of eyes and extending 
to dorsal origin, and a narrow white vertical bar on rear of head be- 
hind eye. Four specimens were studied: USNM 141032, 141033, 
and 147129, from the Marshall Islands and Borneo; and MCZ 33409, 
from Amboina. 


Amphiprion percula (Lacepéde) 
PuaTE 9, Figure C 


Lutjanus percula Lacepéde, Histoire naturelle des poissons, vol. 4, pp. 194, 239, 
240, 1802 (New Britain). 

Amphiprion tunicatus Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 399, pl. 132, fig. 2, 1830 (Vanicolo).—Lesson, Voyage... La 


Coquille . . . , Zoologie, vol. 2, pt. 1, p. 192, pl. 25, fig. 3, 1830 (Port Praslin, 
New Ireland; Doreh, New Guinea). 
Prochilus percula Bleeker, Atlas ichthyologique . . . , vol. 9, pl. 400, fig. 2, 1878. 


Anthias polymnus var. (non Linnaeus) Bloch, Naturgeschichte der auslindischen 
Fische, vol. 6, p. 103, pl. 316, fig. 3, 1792. 

Amphiprion percula Ginther, Journ. Mus. Godeffroy, vol. 15, Andrew Garrett’s 
Fische der Siidsee, pt. 7, pl. 124, fig. A, 1881 (Samoan Islands).—Day, The 
fishes of India . . . , vol. 2, p. 379, pl. 80, fig. 4, 1878 (Andamans).—Montal- 
ban, Bur. Sci. Manila Monogr. 24, p. 14, pl. 2, fig. 2, 1928 (Philippine Islands). 

Actinicola percula Aoyagi, Coral Fishes, Tokyo, pl. 37, fig. 2, 1943 (Kakure- 
Kumanomi); Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, 
p. 175, pl. 9, fig. 2, 1941 (Japan). 

Amphiprion bicolor Castelnau, Proc. Zool. Acclim. Soc. Victoria, p. 92, 1873 
(Port Darwin). 

Actinicola bicolor (Castelnau) Whitley, Mem. Queensland Mus., vol. 9, pt. 3, 
p. 215, pl. 27, fig. 2, 1929 (Port Darwin). 


This is the most characteristically colored species in the genus 
Amphiprion, and is one of the commonest seen associated with the 
sea anemone Discosoma. The second white bar has a forward pro- 
jection under the depressed pectoral fin. No other species of Am- 
phiprion observed by me has the forward edge of the second dark 
bar with a deep concavity. The centers of the dark bars on the 
sides may be pale brown to blackish; all dark bars are black edged. 
I have studied 20 lots, totaling 35 specimens, from the Philippine, 
Solomon, Palawan, Schouten, and Morotai Islands. 


PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


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INDO-PACIFIC FISH GENUS AMPHIPRION—SCHULTZ 193 


Amphiprion laticlavius Cuvier and Valenciennes 
PuatE 9, FigurRE D 


Amphiprion laticlavius Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 394, pl. 132, fig. 1, 18830 (New Guinea). 

Amphiprion bifasciatus (non Bloch) Montalban, Bur. Sci. Manila Monogr. 24, 
p. 15, pl. 3, fig. 1, 1928 (Philippine Islands). 

Amphiprion bifasciatus annamensis Chevy, Travaux |’Inst. Océanogr. Indochine, 
Mem. 4, pt. 1, poissons, p. 99, pl. 39, 1932 (Sud-Annam). 

Prochilus bifasciatus (non Bloch) Bleeker, Atlas ichthyologique . . . , vol. 9, pl. 400, 
fig. 4, 1878. 

Amphiprion polymnus (non Linnaeus) Aoyagi, Coral Fishes, Tokyo, pl. 36, fig. 2, 
1943 (Toaki-Kumanomi); Biogeographica, Trans. Biogeogr. Soc. Japan, 
vol. 4, No. 1, p. 173, pl. 12, fig. 4, 1941 (Japan). 

Amphiprion unimaculatus (non Meuschen) Okada and Ikeda, Biogeographica, 
Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, p. 202, fig. 28, 1939 (Itoman, 
Okinawa). 


This species differs from all others in regard to the second white 
bar, which is represented by an ovate white area on upper half of 
body and on soft dorsal fin, but without any extension on ventral part 
of body. Third white bar on caudal peduncle is lacking. Caudal 
fin is basally and centrally blackish, with posterolateral edges white. 
Anal is submarginally blackish, with distal edge white. One specimen, 
USNM 147128, from the Philippines, was studied. 


Amphiprion chrysogaster Cuvier and Valenciennes 
PuaTE 9, FiaurE EK 


Amphiprion chrysogaster Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 400, 1830 (Ile de France)—Lesson, Voyage . . . La Coquille, ... , 
Zoologie, vol. 2, pt. 1, p. 191, pl. 28, fig. 3, 1830 (fle de France). 

Amphiprion percula (non Lacepéde) Okada and Ikeda, Biogeographica, Trans. 
Biogeogr. Soc. Japan, vol. 3, No. 2, p. 200, pl. 6, fig. 1, 1939 (Riu Kiu Islands), 

Amphiprion trifasciatus Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 595, 1830 (Moluccas). 

Amphiprion fusciventer Bennett, Proc. Comm. Zool. Soc. London, vol 1, p. 165, 
1831 (Mauritius). 

Prochilus bifasciatus (non Bloch) Bleeker, Atlas ichthyologique ... , vol. 9, 
pl. 400, fig. 6, 1878. 


This species, with three broad white bars, has the central part 
of caudal fin black and outer edges white; second white bar is con- 
tinuous on distal part of soft dorsal fin. Seven specimens were 
studied: One each in USNM 61690 from Mauritius, 141034 from the 
Marshall Islands, 147127 from the Philippines; and four MCZ speci- 
mens, three from Zanzibar, Africa, and one, collected by Andrew 
Garrett, from Apiang, Kingsmill Islands. 


194 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Amphiprion polymnus (Linnaeus) 
PuatTE 9, Fiaure J 


Perca polymna Linnaeus, Systema naturae, ed. 10, p. 291, 1758. 

Anthias bifasciatus Bloch, Naturgeschichte der auslindischen Fische, vol. 6, p. 
103, pl. 316, fig. 2, 1792. 

Prochilus bifasciatus Bleeker, Atlas ichthyologique ... , vol. 9, pl. 400, fig. 5, 
1878. 

?Amphiprion ocellaris Cuvier and Valenciennes. Histoire naturelle des poissons, 
vol. 5, p. 399, 1830 (Sumatra). 

?Amphiprion melanurus Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 400, 1830 (Sumatra). 

Lutjanus jourdin Lacepéde, Histoire naturelle des poissons, vol. 4, pp. 191, 235, 
1802 (Amboina). 

Coracinus seu Sciaena unimaculata Meuschen, Zoophylacium Gronovianum ... , 
Pisces, No. 227, 1781 (based on Gronow, 1763). 

Coracinus vittatus Gray, Catalogue of fish collected and described by L. T. Gronow 

. ., British Museum, p. 57, 1854 (based on Gronow’s No. 227). 

Amphiprion intermedius Schlegel and Miiller, Verhandelingen over de Natuurlijke 
Geschiedenis der Nederlandsche Overzeesche Bezittingen ... , Zoologie, 
p. 18, 1839-1841 (reference copied). 


Amphiprion polymnus (Linnaeus) has been confused almost since 
the day 1t was named. Weber and de Beaufort (Fishes of the Indo- 
Australian Archipelago, vol. 8, p. 344, 1940) discuss the confusion 
between polymnus and bicinctus, the former name having been used 
for the species currently called bicinctus. 

In A. polymnus the second white bar continues on to the distal 
part of the soft dorsal, but it lacks the third white bar on the caudal 
peduncle; also, the black coloration of the caudal peduncle continues 
on to the central part of the caudal fin, whereas in A. sebae the caudal 
fin is pale. The anal fin is black basally, with the distal third white. 


EXPLANATION FOR PLATE 9 


A, Amphiprion akallopisos Bleeker, photograph of a color drawing in the Philippine Albatross 
collection; B, 4. perideraion Bleeker, photograph of a color drawing in the Philippine 
Albatross collection; C, 4. percula (Lacepéde), photograph of a color drawing in the 
Philippine Albatross collection; D, A. laticlavius Cuvier and Valenciennes, photograph 
of plate 36, figure 2, in Aoyagi, Coral Fishes, 1943 (= 4. polymnus, non Linnaeus, Aoyagi); 
E, 4. chrysogaster Cuvier and Valenciennes, photograph of plate 6, figure 1, in Okada and 
Ikeda, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, 1939 (=A. percula, non 
Lacepéde, Okada and Ikeda); F, Amphiprion sebae Bleeker, photograph of figure 27 in 
Okada and Ikeda, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, 1939; 
G, A. xanthurus Cuvier and Valenciennes, copy of figure 4 in Jordan and Dickerson, Proc. 
U. S. Nat. Mus., vol. 34, p. 611, 1908, of a specimen from Suva, Fiji; H, A. tricinctus, 
new species, holotype, USNM 152929, from Bikini Atoll, Amen Island, standard length 
75 mm.; 1, 4. mauritiensis, new species, holotype, MCZ 6093, from Mauritius, standard 
length 111.5 mm.; J, 4. polymnus (Linnaeus), photograph of plate 316, figure 2, Bloch, 
Naturgeschichte der auslindischen Fische, vol. 6, 1792 (= Anthias bifasciatus Bloch). 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 9 


Certain species of Amphiprion 
(For explanation, see facing page) 


103, PLATE 10 


PROCEEDINGS, VOL. 


U. S. NATIONAL MUSEUM 


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INDO-PACIFIC FISH GENUS AMPHIPRION—SCHULTZ 195 


I have examined one specimen collected June 16, 1948, by Dr. 
Robert R. Miller in ‘a submerged canoe on a sand bar in Little 
Lagoon, northeast end of Groote Eylandt, Gulf of Carpentaria, 
Australia.’”’ The color when alive was as follows: ‘“The pale bars 
were coral pink, with narrow emerald borders; the dark bars were 
velvet-black.”’ 


Amphiprion tricinctus Schultz and Welander, new species 


PiaTE 9, Figure H 


Amphiprion ephippium (non Bloch) var. chrysopterus (non Cuvier and Valen- 
ciennes) Giinther, Journ. Mus. Godeffroy, vol. 15, Andrew Garrett’s Fische 
der Siidsee, pt. 7, pp. 224-225, pl. 122, fig. C, 1881 (Kingsmill Islands). 


Holotype—USNM_ 152929, Bikini Atoll, Amen Island, lagoon, 
August 21, 1947, University of Washington, Staff of Applied Fisheries 
Laboratory, standard length 75 mm. 

Description.—Dorsal fin rays X,17; anal II,14; pectoral 11,17,i to 
ii,16,ii; pelvics I,5; branched caudal fin rays 8+7; vertical scale rows 
from upper edge of gill opening to base of caudal fin 54, scales between 
lateral line and base of first soft dorsal ray 5, and between lateral 
line and anal origin 20; pores in lateral line 36; predorsal scales 19 or 
20; gill rakers 5-+1-+13. 

Detailed measurements were made on the holotype and these data 
are expressed in thousandths of the standard length, 75 mm.: Greatest 
depth 560; length of head 272; snout 99; eye 95; least preorbital 35; 
length from snout tip to rear edge of maxillary 101; postorbital length 
of head 147; least width of interorbital space 100; least depth of 
caudal peduncle 160; length of caudal peduncle from base of last anal 
ray to midbase of caudal fin 192; length of longest ray of pectoral 287, 
pelvic 313, upper caudal fin 307, lower lobe of caudal fin 300, spiny 
dorsal 160; length of next to last dorsal spine 160; width of white 
part of first pale bar at level of lateral line 73, second 40, last (pe- 
duncular) 20. 

Depth of body 1.7, head 3.4, both in standard length. Snout 3.2; 
eye 3.1; least preorbital distance 8.0; upper jaw 2.8; postorbital part 
of head 2.0; least interorbital space 3.0; least depth of caudal peduncle 
1.8; length of pectoral fin 1.0, pelvic 0.9, second dorsal spine 23, 
upper caudal rays 0.9 to 1.0; all in length of the head. Least depth 
of caudal peduncle in its length 1.2. Angle of upper profile of head 
with lengthwise axis of body about 50°; profile of head convex. 

Teeth in both jaws in a single row, nearly conical, a little com- 
pressed forward, pointed; interorbital space scaled forward to a line 
between middle of pupils; 4 or 5 rows of scales on cheeks; gill cover with 
a few scales; scales occur part way out on all median fins; preorbital 
with 3 spines; suborbital with 10 to 12 smaller spines. 


196 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Color in alcohol.—Background coloration of body from about fifth 
dorsal spine posteriorly and dorsally to midlengthwise axis of body 
blackish, the anteroventral part of body pale light brown becoming 
paler ventrally; spiny dorsal dark brown; soft dorsal black; caudal fin 
black, except edged with white posteriorly; pectoral and pelvic fins 
pale or very light tan; anal pale or light tan, distally edged with a black 
line; first white bar begins a little in front of dorsal origin and just 
behind eye, ending on lower edge of subopercle; second white bar 
begins on last dorsal spine and base of first soft ray in lower third of 
fin, extends ventrally, meeting its fellow in narrow space between 
anal origin and anus; third white bar, about half width of second, or 
narrower than width of pupil, crosses caudal peduncle just in front of 
caudal fin base. 

Remarks.—This new species may be recognized by the narrowness 
of the three white bars, especially the second and third, and by the 
black caudal fin narrowly edged with white posteriorly. It is sep- 
arated from all other species of Amphriprion by the key. 

Named ¢ricinctus in reference to the three white bars. 


Amphiprion mauritiensis Schultz, new species 


Puate 9, Ficure I 


Holotype-—MCZ 6093, Mauritius, collected by Nicolas Pike, stand- 
ard length 111.5 mm. 

Paratypes.—Bearing same data as holotype: Out of MCZ 6093, 4 
specimens, 67 to 96 mm.; MCZ 5801, 2 specimens, 98 and 104.5 mm.; 
MCZ 5800, 1 specimen, 101.5 mm.; MCZ 5802, 2 specimens, 83.5 and 
114 mm. 

Description.—Dorsal rays X,17 or 18, one with XI,16, usually 
X,17; anal III,14, one with III,13; pectoral rays 20 or 21; pelvics 
1,5; branched caudal fin rays 8+7; vertical scale rows from upper 
edge of gill opening to base of caudal fin about 55 to 58; scales between 
lateral line and base of first soft dorsal ray 5; between lateral line and 
anal origin 17 to 19; pores in lateral line 35 to 42; predorsal scales 
about 10 or 11; gill rakers 5+1+13. 

Detailed measurements were made on the holotype and two para- 
types, and these data are expressed in thousandths of the standard 
length, first for the holotype then, in parentheses, for the paratypes: 
Standard length in mm. 111.5 (87.5; 104.5). Greatest depth 485 
(490; 535); length of head 297 (286; 310); snout 90 (97; 96); eye 89 
(86; 86); least preorbital width 31 (38; 28); distance from snout tip 
to rear edge of maxillary 91 (114; 112); least width of interorbital 
space 74 (91; 81); least depth of caudal peduncle 144 (157; 151); 
length of caudal peduncle 166 (153; 148). Length of longest ray of 


INDO-PACIFIC FISH GENUS AMPHIPRION—SCHULTZ 197 


pectoral 279 (269; 280); pelvic 291 (257; 296); upper caudal lobe 270 
(280; 296); lower caudal lobe 265 (263; 277); spiny dorsal 135 (132; 
124). Length of next to last dorsal spine 99 (86; 86). Width of 
first white bar at level of lateral line 81 (80; 67); second 71 (74; 57); 
third (peduncular) 48 (63; 48). Width of second black bar at level 
of lateral line 323 (831; 344); third 300 (292; 344). 

Depth of body 1.8 to 2.1, head 3.2 to 3.5, both in standard length. 
Snout 2.9 to 3.2; eye 3.3 to 3.6; least preorbital 7.5 to 10.1; upper 
jaw 2.5 to 3.2; least interorbital 3.1 to 3.9; least depth of caudal 
peduncle 1.9 to 2.0, length of pectoral 1.1, pelvic 1.0 to 1.1, third 
dorsal spine 2.1 to 2.5; all in length of head. Least depth of caudal 
peduncle in its length 0.95 to 1.2. Width of second white bar at 
level of lateral line in width of third black bar at level of lateral line 
3.5, third white bar 4.0 to 7.0. 

Profile of head convex. Teeth uniserial, nearly conical, a little 
compressed forward, pointed; interorbital space scaled forward to a 
line between rear of eyes; 4 to 6 rows of scales on cheek; gill cover 
with a few scales; scales occur part way out on all median fins; pre- 
orbital with 1 to 3 spines; suborbital with about a dozen spines. 

Color in alcohol.—Background coloration dark brownish or blackish, 
except that underside of head and ventrally below a line from pectorals 
to anus is white or pale; three white bars, one behind eye, second from 
origin of soft dorsal to anus, third on caudal peduncle; dorsal fin 
black, with distal edge of soft dorsal white; anal fin black or pale, if 
black, distal edge is white; pelvics pale; pectoral pale, except dusky 
basally on some specimens; central area of caudal fin blackish, edges 
white, with edge widest distally; gill membranes appear to be white. 

Remarks.—This new species is most closely related to A. tricinctus 
but differs in having wider white bars, as compared in the key. 
Named mauritiensis in reference to the locality where it was collected, 


Amphiprion sebae Bleeker 


PuaTE 9, Figure F 


Amphiprion sebae Bleeker, Nat. Tijdschr. Nederl.-Indié, vol. 4, p. 478, 1853; 
Atlas ichthyologique ..., vol. 9, pl. 400, fig. 9, 1878.—Day, Fishes of 
India ..., vol. 2, p. 378, pl. 80, fig. 8, 1878 (Andamans).—Okada and 
Ikeda, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 2, p. 200, 
fig. 27, 1939 (Isigaki and Irimote Islands). 


This species, in which the second white color bar continues distally 
on the soft dorsal fin, is very much like A. polymnus, but differs in 
having a white bar on the caudal peduncle and a white caudal fin. 
Three specimens were studied: USNM 45169, from the Seychelles, 
and 133830, from the Paumotu Islands; and MCZ 33410, from Sumatra. 


198 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Amphiprion xanthurus Cuvier and Valenciennes 


PLaTE 9, FIGURE G 


Amphiprion xanthurus Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 402, 1830 (Ile de France). 

Amphiprion clarckit Cuvier and Valenciennes, Histoire naturelle des poissons, 

>& vol. 9, p. 504, 1833 (emended spelling on Bennett). 

Anthias clarkii Bennett, A selection from . . . fishes found on the coast of Ceylon, 
London, ed. 2, p. 29, pl. 29, 1834 (Ceylon). 

Sparus milli Bory de Saint-Vincent, Dictionnaire classique d’histoire naturelle, 
vol. 17, p. 130, pl. 113, fig. 2, 1831 (China Sea). 

Prochilus polymnus (non Linnaeus) Bleeker, Atlas ichthyologique, vol. 9, pl. 400, 
figs. 7, 8, 1878. 

Anthias polymna (non Linnaeus) Bloch, Naturgeschichte der auslindischen 
Fische, vol. 9, p. 89, pl. 316, fig. 1, 1792. 

Amphiprion bicinctus (non Rippell) Aoyagi, Coral Fishes, Tokyo, pl. 37, fig. 1, 
1943 (Kumanomi).—Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 4, 
No. 1, p. 169, pl. 9, fig. 1, 1941 (Japan). 

Amphiprion polymnus (non Linnaeus) Okada and Ikeda, Biogeographica, Trans. 
Biogeogr. Soc. Japan, vol. 3, No. 2, p. 204, fig. 30, 1989 (Riu Kiu Islands) .— 
Montalban, Bur. Sci. Manila Monogr. 24, p. 10, pl. 1, fig. 1, 1928 (Philippine 
Islands). 

Amphiprion chrysopterus Cuvier and Valenciennes, Histoire naturelle des poissons, 
vol. 5, p. 401, 1830 (no locality given).—Jordan and Dickerson, Proc. U. 8. 
Nat. Mus., vol. 34, p. 612, fig. 4, 1908 (Suva, Fiji). 

Amphiprion japonicus Temminck and Schlegel, Fauna Japonica ... , Pisces, 
p. 66, 1843 (Japan). 

Amphiprion chrysargyrus Richardson, Rep. Meetings British Assoc. Adv. Sci., 
vol. 15 (1845), p. 254, 1846 (Seas of China and Japan). 

Amphiprion boholensis Cartier, Verh. phys. med. Wirzburg, new ser., vol. 5, 
p. 96, 1874 (Bohol). 

Amphiprion melanostolus Richardson, Ann. Mag. Nat. Hist., vol. 9, p. 390, 1842 
(Depuch Island). 

?Amphiprion de bojer Lienard, Treizisme Rapp. Ann. Soc. Hist. Nat. Maurice, 
p. 68, 1843 (reference copied). 

Amphiprion snyderi Ishikawa, Proc. Nat. Hist. Tokyo Mus., vol. 1, No. 1, p. 11, 
pl. 5, 1904 (Bonin Island). 


I have examined 70 specimens (in 45 lots) of zanthurus, 22 of which 
are in the Museum of Comparative Zoology, Harvard University, 
and the others in the U.S. National Museum, and I find some variation 
in color pattern. Among these specimens 12 had black dorsal, anal, 
and pelvic fins; 3 had these three fins dusky; 10 had the dorsal and the 
anal black or dusky, with pelvics pale; 36 had the dorsal black and 
both anal and pelvics pale; 9 had dorsal, anal, and pelvics pale, except 
in some specimens the pelvic fins were dark edged. Of these 70 
specimens 42 were from the Philippines, 4 were from Japan, 2 from 
the China coast, 18 from Zanzibar, 1 from the Gilbert Islands, and 
3 from Kingsmill Islands. 


INDO-PACIFIC FISH GENUS AMPHIPRION—SCHULTZ 199 
Amphiprion bicinctus Riippell 
PuaTe 10, Figure A 


Amphiprion bicinctus Riippell, Atlas zu der Reise im nérdlichen Afrika, p. 139, 
pl. 35, fig. 1, 1828 (Red Sea). 

Amphiprion papuensis Macleay, Proc. Linn. Soc. New South Wales, vol. 8, No. 
2, p. 271, 1883 (New Guinea).—Whitley, Mem. Queensland Mus., vol. 9, 
pt. 3, p. 210, pl. 27, fig. 1, 1929 (D’Entrecasteaux Group, New Guinea, on 
holotype). 

Amphiprion arion De Vis, Proc. Linnean Soc. New South Wales, vol. 8, p. 450, 
1884 (South Seas). 


This species has the first two white bars but lacks the peduncular 
bar; the anterior edges of the pelvics are black and the caudal fin is 
pale. Six lots containing seven specimens were studied: USNM 
61679, from Suva, Fiji; USNM 141030 and 141031, from the Marshall 
Islands; uncataloged specimens, one from the Paumotu and two with- 
out data; also one from Bikini in the University of Washington 
collection. 

Among these specimens, four adults, 82 to 99 mm. in standard length, 
have a pale anal fin, whereas two, 26 and 27 mm. long, have a black 
anal, and one, 25 mm. long, has a dusky anal fin. The caudal peduncle 
varies from pale dusky to black or brownish. The key gives the 
essential color pattern of this species. 


Amphiprion ephippium (Bloch) 
PuaTe 10, Ficures B, C 


Lutjanus ephippium Bloch, Naturgeschichte der auslandischen Fische, vol. 4, 
p. 121, 1790 (reference not seen) ; Ichthyologie, ou histoire naturelle, générale 
et particuliére des poissons... , vol. 7, p. 98, pl. 250, fig. 2, 1797 (East Indies). 

Amphiprion ephippium Day, Fishes of India . . . , vol. 2, p. 378, pl. 80, fig. 1, 
1878. 

Amphiprion monofasciatus Thiolliére in Montrouzier, Suite de la faune de I’Ile 
de Woodlark ou Moiou, Ichthyologie, Ann. Sci., Physic. Nat. Agr. Indust., 
Lyon, vol. 8, p. 476, 1856 (Woodlark Island). 

Amphiprion tricolor Giinther, Catalogue of the fishes in the British Museum, 
vol. 4, p. 8, 1862 (Port Essington; South Australia). 

Amphiprion riippeli Castelnau, Proc. Zool. Acclim. Soc. Victoria, vol. 2, p. 91, 
1873 (Port Darwin). 


Amphiprion frenatus (non Brevoort) Day, Fishes of India . . . , vol. 2, p. 378, 
pl. 80, fig. 2, 1878. 

Prochilos ephippium Bleeker, Atlas ichthyologique ..., vol. 9, pl. 401, figs. 
1, 9, 1878. 


Amphiprion rubrocinctus Richardson, Ann. Mag. Nat. Hist., vol. 9, p. 391, 1842 
(Depuch Island; probably young). 
The adults of this species usually have a plain blackish body and 
head, with a single white color bar on the head, but on a few speci- 
mens, the largest in the series, this first white bar is lacking or nearly 


200 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


so. The smallest specimens among 57 collected by Dr. Robert R. 
Miller from Woods Inlet west of Darwin, Australia, have a variable 
color pattern of two or three white bars. One specimen, 20 mm. in 
standard length, has only one white bar on the head; 9, from 20 to 
31 mm., have two white color bars, the first and second; 7, from 19.5 
to 28 mm., have the first and second white bars distinct and the third 
indistinct, sometimes represented by a white spot on dorsal edge of 
caudal peduncle; 10, from 17 to 25 mm., have all three white color 
bars distinct. I presume that the second and third white bars dis- 
appear with increase in size, since a 35-mm. specimen in this lot looks 
like all the larger specimens with only one white bar on the head. 


Amphiprion frenatus Brevoort 
PuaTE 10, Figure D 


Amphiprion frenatus Brevoort, U. S. Japan Exped. Nat. Hist., Washington, 
vol. 2, p. 263, pl. 6, fig. 4, 1856 (Lew Chew [Okinawa]).—Jordan and Snyder, 
Proc. U. S. Nat. Mus., vol. 24, p. 597, 1902.—Montalban, Bur. Sci. Manila 
Monogr. 24, p. 12, pl. 2, fig. 1, 1928 (Philippine Islands).—Aoyagi, Biogeo- 
graphica, Trans. Biogeogr. Soc. Japan, vol. 4, No. 1, p. 167, 1941 (Japan).— 
Okada and Ikeda, Biogeographica, Trans. Biogeogr. Soc. Japan, vol. 3, No. 
2, p. 203, fig. 29, 1939 (Riu Kiu Islands). 

Prochilus polylepis Bleeker, Versl. Akad. Wet. Amsterdam, ser. 2, vol. 11, p. 135, 
1877; Atlas ichthyologique . . . , vol. 9, pl. 401, fig. 6, 1878. 

Amphiprion polymnus (non Linnaeus) Montalban, Bur. Sci. Manila Monogr. 
24, p. 10, pl. 1, fig. 1, 1928 (Philippines). 

I have studied 72 specimens from the Philippines, Okinawa, and 
Japan referable to this species. In alcohol two color phases are evi- 
dent, the usual one (53 specimens, ranging from 37 to 100 mm. in 
standard length) has a plain blackish body behind head, whereas 15, 
ranging from 39 to 60 mm., have 3 lengthwise pale bands on side of 
body, and 4 others are intermediate in regard to coloration. These 
pale bands at best are not very distinct. This species is character- 
ized by having about twice as many of the specimens with IX dorsal 
spines as with X; this latter figure is the usual number of dorsal 
spines in the other species of Amphiprion. 

Of 78 specimens in 54 lots studied, 76 were from the Philippine 
Islands and one each from Japan and Okinawa. The specimen from 
Okinawa (the type locality of frenatus), USNM 71702, Naha, Okinawa, 
Albatross, 69 mm. in standard length, is herewith designated as neo- 
type of Amphiprion frenatus Brevoort since the type is not in existence. 


INDO-PACIFIC FISH GENUS AMPHIPRION—SCHULTZ 201 


Amphiprion melanopus Bleeker 
Puate 10, Ficure E 


Amphiprion melanopus Bleeker, Nat. Tijdschr. Nederl. Indié, vol. 3, p. 561, 1852. 

Prochilus melanopus Bleeker, Atlas ichthyologique ... , vol. 9, pl. 401, fig. 7, 
1878. 

Prochilus macrostomus Bleeker, Atlas ichthyologique . . . , vol. 9, pl. 401, fig. 5, 
1878. 

Amphiprion ephippium (non Bloch) Giinther, Journ. Mus. Godeffroy, vol. 15, 
pt. 7, pl. 122, fig. D (var. melanopus on p. 225), 1881. 

Amphiprion mccullochi Whitley, Mem. Queensland Mus., vol. 9, pt. 3, p. 213, 
1929 (Lord Howe Island). 

Amphiprion macrostoma (non Bleeker) Chevy, Travaux Inst. Oceanogr. Indo- 
Chine, Mem. 4, pt. 1, Poissons, p. 102, pl. 40, 1932 (Annam). 


This species is characterized by the single white bar on the head, 
plain black body, pale soft dorsal, caudal, and pectoral, and black 
pelvics and anal fins. 

Of 18 specimens in 7 lots studied, 14 were from the Marshall 
Islands, 3 from Apia, Samoa, and 1 from Paumotu Island. 


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SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington : 1954 No. 3324 


MARINE POLYCHAETE WORMS FROM POINT BARROW, 
ALASKA, WITH ADDITIONAL RECORDS FROM THE 
NORTH ATLANTIC AND NORTH PACIFIC 


By Marian H. PETTIBONE 


This report of the Arctic polychaetes found in the region of Point 
Barrow, Alaska, is based on material collected during 1948, 1949, and 
1950 by G. E. MacGinitie, of the Arctic Research Laboratory. Speci- 
mens were obtained from Eluitkak Pass, Elson Lagoon, near Point 
Barrow, were washed ashore at the Point Barrow base, and were 
dredged within 16 miles offshore at Point Barrow base in depths of 
1.7 to 123.5 fathoms on bottoms of mud, stones, gravel, rocks, in 
masses of worm tubes, and various combinations of these. Addi- 
tional specimens were collected from fish traps, from screen traps 
lowered through holes made in the ice, and from plankton hauls, 
some of which were made through holes in the ice. The latter collec- 
tions are of particular interest in that they include specimens showing 
sexual epitokous stages of some of the syllids. Considerable care 
was taken in going over the miscellaneous material and separating 
polychaetes, as evidenced by the large collection and the presence 
in it of many small specimens of the young of larger species as well 
as small species which are often overlooked. There are a great num- 
ber of small syllids, phyllodocids, hesionids, and terebellids. Great 
care also was taken in the preservation of the specimens. Some 
color notes were included and some color photographs were taken, and 
these added considerably to the value of the collection. 

The material was worked on by the writer at the U. S. National 
Museum, where the collection is deposited. The facilities of the Mu- 
seum, including laboratory accommodations, the library, and the vast 

261112—54——_1 203 


204 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


polychaete collections, which include much type material, were placed 
at the writer’s disposal. Some previously unworked material was 
examined for comparative purposes, and the results of some of these 
related studies are included here. This material is chiefly from collec- 
tions made by the following: R. A. Bartlett in the Canadian Arctic, 
Greenland, and Labrador from 1927 to 1942; W. H. Dall in Arctic 
Alaska, the Bering Sea, and southeastern Alaska from 1871 to 1880; 
the Blue Dolphin expeditions to Labrador and Newfoundland from 
1949 to 1951 under the command of D. C. Nutt; the U. S. Fish Com- 
mission in dredgings off the east coast of North America, from which 
many specimens had been identified and recorded by A. E. Verrill; 
and by the writer in the Straits of Juan de Fuca, in Washington and 
Puget Sounds, Washington, chiefly during the summers from 1936 to 
1940, and from the region of Woods Hole, Massachusetts, in the 
summers of 1950 and 1951. For the last three collections mentioned, 
only summaries of the data are given for species common to the 
Point Barrow region. More complete data are to be published sepa- 
rately and the summaries are included here only to make the dis- 
tributional data of the Point Barrow species more complete. 

The number of polychaetes previously recorded from Arctic Alaska 
is small indeed. The collection obtained by the International Polar 
Expedition to Point Barrow from 1881 to 1883 included only 17 poly- 
chaetes (Murdoch, 1885, p. 152). This collection, which was depos- 
ited in the U. S. National Museum, was examined by the writer and 
is referred to in this report. A few additional records in scattered 
papers have added to the list of polychaetes of Arctic Alaska. 

For each of the species, a rather full but by no means complete 
synonymy is given, chiefly bringing together references to the original 
and additional descriptions and scattered distributional records. To 
facilitate identification, keys to the families, genera, and species are 
given, as well as synopses of the families and brief descriptions of the 
species with size ranges and color notes. An explanatory key to the 
lettering of the diagnostic features in the figures is given on page 210. 

This study was aided by a contract between the Office of Naval 
Research, Department of the Navy, and Johns Hopkins University 
(Project No. NR 162 911, Contract and Task Order No. N6onr 
243-16). 

The writer acknowledges her appreciation to Prof. and Mrs. G. E. 
MacGinitie for their cooperation and help on this project; to the au- 
thorities of the U. S. National Museum for allowing her to make use 
of the facilities of the Institution, especially to Dr. Waldo L. Schmitt 
and Dr. Fenner A. Chace, Jr., for their valuable aid, suggestions, and 
patience; and to Mr. and Mrs. Cyril Berkeley, of Nanaimo, British 
Columbia, for their loan of specimens and their valuable suggestions. 


MARINE POLYCHAETE WORMS—PETTIBONE 205 


The distribution of the Point Barrow polychaetes is summarized 
in table 1. 

The collections include 3,270 specimens representing 88 species and 
26 families of Polychaeta. Some of the species were exceedingly 
common; others were represented only by single or few specimens. 
The Polynoidae, Syllidae, and Terebellidae are the most abundant 
both as to numbers of species (11 each, or 37 percent) and specimens 
(66 percent). The Phyllodocidae and Sabellidae have 7 species each. 
Eleven families are represented by a single species each. 

During 1948, because of the ice floes all summer, there was no 
heavy surf and practically nothing was washed up on the beach. 
During 1949, specimens were washed ashore on 17 different days. 
In 1950, specimens were collected from the beach on only one day. 
Altogether, 287 specimens of polychaetes were washed ashore, repre- 
senting 20 species and 11 families. Two species, Hunoé clark (2 
specimens) and Travisia carnea (12 specimens), were obtained in this 
manner only. Four species were dredged occasionally but were 
washed ashore in much larger numbers, namely: Antinoé sarsi (58 
specimens), Melaenis lovént (65 specimens), Arenicola glacialis (54 
specimens), and Brada villosa (24 specimens). Additional species 
were commonly obtained in the dredge but were washed ashore in 
small numbers. 

On three occasions in 1950 when 80-foot vertical plankton hauls 
were taken through holes in the ice, the sexual stages of three of the 
syllids were found—Autolytus fallax, Syllis cornuta, and Syllis fasciata. 

A station 12.1 miles from shore, 123.5 fathoms, August 17, 1949, 
produced literally bushels of worm tubes. The tubes of the terebellid 
Pista maculata made up the greater part of the mass. 

Of the 88 Point Barrow species, 75 (85.2%) are common to the 
Arctic, Atlantic, and Pacific; 5 (5.7%) are common only to the Arctic 
and Pacific (Gattyana ciliata, Eusyllis magnifica, Pionosyllis compacta, 
Glycinde wiréni, and Idanthyrsus armatus); 3 (3.4%) are common to 
the Arctic and Atlantic (Autolytus fallar, Eumida minuta, Travisia 
carnea); 4 (4.5%) are confined to the Arctic (Hunoé clarki, Nerinides 
sp., Arenicola glacialis, Ampharete vega); and 1 (1.1%) is bipolar, 
known only from the Arctic and Antarctic (Ammotrypane breviata). 

The Point Barrow records help to complete the circumpolarity 
records for 58 circumpolar or almost circumpolar species, some of 
which previously had been known only from the Siberian and Canadian 
Arctic, Greenland, and Spitsbergen. The range has been extended 
for 28 species, 6 of which have been extended by combining Atlantic 
and Pacific specific names. 


PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


206 


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VOL. 103 


PROCEEDINGS OF THE NATIONAL MUSEUM 


208 


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209 


MARINE POLYCHAETE WORMS—PETTIBONE 


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(suommoods sGp) AVAIITAAAAD, 


210 PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 103 


Explanation of symbols on figures 


Roman numerals indicate body segments. 


a, anus 

aBr, anal branchia 

aC, anal cirrus 

aCo, anal cone 

aCy, anal cylinder 

ak, anal knob 

aLi, anal ligule 

abd, abdominal region 

ac, aciculum (heavy spine buried in 
parapodial ramus) 

acH, acicular genital hook 

acL, acicular lobe 

ach, achaetous lobe 

ai, aileron of jaw 

an, antenna 

antLa, anterior or presetal lamella 

anta, antanal achaetous segment 

ba, barbule 

bl, blade of seta (terminal or end 
piece of composite or jointed seta) 

br, branchia, 

brF, branchial filament 

brLo, branchial lobe 

brP, brood pouch 

buS, buccal segment 

cP, cephalic peak 

cCre, cephalic crest 

cPl, cephalic plate 

cR, cephalic ridge 

caPl, calcareous plate 

co, collarette 

eph, cirrophore (base of a tentacular 
cirrus or a dorsal cirrus) 

cr, crotchet 

cre, crest 

dC, dorsal cirrus 

dL, dorsal lobe 

dN, dorsal notch 

dTu, dorsal tubercle (lobe correspond- 
ing to elytrophore on non-elytra- 
bearing segment) 

deR, denticled rim 

Dor, dorsal 

el, elytron or scale 

elph, elytrophore (lobe bearing ely- 
tron) 

ey, eye-spot 

fTu, facial tubercle 

frAn, frontal antenna 


frH, frontal horn 

j, jaw 

1An, lateral antenna 

1Gr, lateral groove 

Lo, lateral lobe 

1N, lateral notch 

10Or, lateral sensory organ 

liPl, limbate plate 

loL, lower lip 

mAn, median antenna 

mLo, median lobe 

maxR, maxillary or distal ring 
mo, mouth 

nPa, nephridial papilla 

ne, neuropodium or ventral ramus 
neC, neuropodial cirrus 

neLi, neuropodial ligule 

neS, neuroseta (seta of neuropodium) 
neTo, neuropodial torus 

no, notopodium or dorsal ramus 
noLi, notopodial ligule 

nos, notoseta (seta of notopodium) 
noPi, notopodial pinnule 

noTo, notopodial torus 

nuEp, nuchal epaulette 

nuF, nuchal fold, lobe or collar 
nuG, nuchal groove 

nuH, nuchal hook 

nuO, nuchal organ 

nuPa, nuchal papilla 

oR, oral or basal ring 

oT, oral tentacles 

ocT, occipital tubercle 

op, operculum 

opPe, opercular peduncle 

Pp, palp 

pM, palmar membrane 

pa, papilla 

pal, paleae 

parF, parapodial flange 

para, paragnaths or horny denticles 
parath, parathoracic segment 
poL, postsetal or posterior lobe 
poLa, postsetal or posterior lamella 
pr, prostomium 

preL, presetal or anterior lobe 
pro, proboscis 

pyF, pygidial or anal funnel 

pyPl, pygidial plate 


MARINE POLYCHAETE WORMS—PETTIBONE 211 


8, seta 

scH, scaphal hook 

set, setigerous segment or setiger 

setLo, setigerous lobe 

sh, shaft of seta (basal part of com- 
posite or jointed seta) 

spH, special hook 

tC, tentacular cirrus 

tF, tentacular filament 

tM, tentacular membrane 

tS, tentacular segment or peristomium 
(anterior modified segments around 
mouth) 

th, thoracic region 

thM, thoracic membrane 

to, tooth 

tor, torus (low, inflated parapodial 
lobe) 


uL, upper lip 

un, uncinus (very small and short, 
flattened setal hooks) 

unPi, uncinigerous pinnule (flattened 
parapodial ramus bearing numerous 
uncini or small setal hooks) 

unTo, uncinigerous torus (low, in- 
flated parapodial ramus _ bearing 
numerous uncini or small setal 
hooks) 

vC, ventral cirrus 

vF, ventral faecal groove 

vL, ventral lobe 

vPa, ventral papilla 

vsh, ventral shield 

Ven, ventral 

w, basal crenulate wing 


Order POLYCHAETA 


Key to the families of Polychaeta from Point Barrow 


. Dorsal surface more or less completely covered by paired overlapping scales 

SiC Vbrin A 2ON Mt Yh. | tet ee eee eS ee eet eS 5 
Dorsal surface not covered by elytra..............--.--..------------- 2 
. Anterior end more or less concealed by feathery tentacles (fig. 38, a), chiti- 
nous golden setae forming an operculum (figs. 34, n; 35, d), long setae 
directed forward forming a cephalic cage, or long filamentous outgrowths 


(fig. 36, f). With more or less permanent tubes-_-_-.-....---...----.- 6 
Ii CMC ETINIRG hb lita re teeeee! sawn eo beeen tases 2d 3 

_ body thickly papillated, grub-shaped. oo4i0.20.% 2. Jos24-e-e+-n22k--~ 21 
Bady airoothjor finely, granulated... 4,522. 222.222) LA beet stds 4 


. Prostomium with conspicuous antennae, with or without fleshy palps (figs. 
26-30), or prostomium or an anterior segment with pair of long, prehensile, 


tenincular pulps (fig. 32,4) t1o.<eo- ot at etetic be Secs ays Seta e 13 
Prostomium without palps and without conspicuous antennae (fig. 32, a) or 
with 2 pairs minute antennae (figs. 30, j; 31, a, e)------------------- 18 

. Elytra on all segments of posterior region (at least from segment 29 on). 
Marsal CrThwacking |... <<. ote oo cecalcard datenioe Sigalionidae (p. 228) 
Elytra not on all segments of posterior region. With paired dorsal cirri on 
segments without elytra (fig. 26, a)._-.------------- Polynoidae (p. 213) 

. Anterior end with branchial plume (fig. 38, a)_.--.--------------------- if 
Anterioriend, without, branchial plume.=s-=< 22-2 -222-)5-24)-25 4-2 24--2- 8 


. Branchial filaments without operculum. Without thoracic membrane. 
Tube gelatinous, membranous, or horny, covered or not with mud, sand, 
Braver, and deprig= = 9-2-2 as eee Sabellidae (p. 332) 

Usually 1 or 2 opercula on modified branchial filaments (fig. 39, uw). Usually 
with thoracic membrane (fig. 39, uw). Tube calcareous__Serpulidae (p. 342) 

. Anterior end with chitinous, golden setae forming an operculum (figs. 34, 
n; 35, d). In free, conical, sandy tubes or concreted sandy tubes attached 
to Tocks, shells) ete... == pellnee ot Leth vernee teed sp no eet S-- = 12 

Anterior end without opercular setae. 


212 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


9. 


10. 


11. 


12. 


13. 


14. 


15. 


16. 


1 Ls 


18. 


19: 


Setae of anterior segments directed anteriorly, forming a cephalic cage. 
Body covered with papillae and often with adherent sand grains; may be 
encased in thick mucoid sheath._______________- Flabelligeridae (p. 288) 

Anterior end without cephalic cage but with long, filamentous outgrowths 
(may have 2 groups of paleal setae directed forward; fig. 36, a, e, f). Body 
without papilacsi!? (Cuvee 2255-22 =- 5s eee 10 

Anterior end with long, filamentous outgrowths and with similar outgrowths 
continued along sides of body on few to many segments (fig., 33, a). Seg- 
ments similar, body not divided into 2 distinct regions. 

Cirratulidae (p. 285) 

Long, filamentous outgrowths concentrated on anterior region only (figs. 36, 
37). Body'with 2 distinct’regions__- + S08_°0°2') 203 See ee 11 

With numerous filiform oral tentacles retractile in mouth (fig. 36, e). Ante- 
rior region with or without 2 groups of paleal setae directed forward (fig. 
SO, G20) oe 2 Seen een: Mee Oe a kee Ampharetidae (p. 314) 

With numerous, grooved, oral tentacles not retractile in mouth (fig. 36, f, g, 
t). Anterior region without paleal setae________-__- Terebellidae (p. 318) 

Opercular setae forming 1-3 concentric rows (fig. 34, n-g). Tube of con- 
creted sand or fine gravel, attached____________-- Sabellariidae (p. 307) 

Opercular setae forming 2 bundles in horizontal rows, directed obliquely 
anteriorly (fig. 35, c, d). Tube free, conical, rigid, open at both ends, 


formed of single layer of sand grains_-.---------- Pectinariidae (p. 310) 
Prostomium with 2—5 conspicuous antennae. With 1-8 pairs tentacular cirri 
(figs! 27-39) UO Ee Ses Se) EE Sent I 2 See 14 


Prostomium without conspicuous antennae. Without tentacular cirri. With 
pair of tentaculiform, longitudinally-grooved palps (deciduous, may be 
Missing: HES" 32,'t; 30, a) 2224 TS. ONO) Se ee eee 17 

Dorsal cirri flattened, foliaceous, sometimes overlapping, or more or less 
globular (fig. 27). Prostomium with 2 eyes, 4-5 antennae, without 


Palus-!222>s<2+-= UR SUBSE fare. Se eee Phyllodocidae (p. 231) 
Dorsal cirri otherwise. Prostomium with 4-6 eyes, 2-3 antennae, 2 palps 
(may be reduced and fused in -Syllidae) __+_/2_~_- 4-22 Vee eee 15 
With parapodial ligules, rounded, conical or straplike lobes (fig. 30, b, c). 
Worsalcirr short, cirriform: se Se2s Se eee ee Nereidae (p. 263) 
Without parapodial ligules. Dorsal cirri filiform, may be more or less articu- 
lated (or small and subulate in some Syllidae) _.__._._.__________-_-- 16 
Palps ‘biartieulgte ig.’ 28;a) 222 22022 ee ae 2008 Hesionidae (p. 239) 


Palps not biarticulate (may be reduced and fused; figs. 28, c; 29, a). 
Syllidae (p. 240) 
Parapodia with postsetal lamellae more or less developed (fig. 32, g). Branch- 
iae usually paired, dorsal, ligulate or cirriform (rarely pinnate, rarely 
Inekinpy2 G8 £4 2+ ce eee ee EE OR ee Oe Spionidae (p. 280) 
Parapodia with lobes scarcely projecting, without postsetal lamellae. Branch- 
iae simple, filamentous, contractile, inserted above notopodia on few to 


many segiiients (fig) 33) d) ULL Lo52e JuoLsie Be Cirratulidae (p. 285) 
Prostomium with 2 pairs minute antennae (figs. 30, 7; 31, a@)_-_--_------ 19 
Prostomium without-antennae.~.—..._ == ==. = ae ee See 22 


Prostomium subquadrate, with small antennae on anterior part (fig. 30, 7). 
With anterior and posterior parapodial lamellae more or less developed 
(fig SO %n)> Sel ASIN ob ies BONN UU A Es ey Fe Nephtyidae (p. 266) 

Prostomium conical, pointed, annulated, with tiny terminal antennae (fig. 
31, a, e). . Without parapodial lamellae... —. 22 _ Suede 2a eee 20 


MARINE POLYCHAETE WORMS—PETTIBONE 213 


20. Segments bi- or triannulate. Parapodia of single form throughout length of 


body, either only uniramous or only biramous------- Glyceridae (p. 272) 
Segments uniannulate. Body divided into 2 regions—anterior uniramous, 
posterior biramous (fig; 31, f; 9) ==25==2222--=-22022 Goniadidae (p. 274) 

21. Posterior part of body with 2 horny ventral plates with radiating bundles of 
Bette (hee oO) eee ee ee, SIR) Tate SOR Te 290 Sternaspidae (p. 309) 
Posterior part of body without horny plates_-_--_--- Flabelligeridae (p. 288) 


22. Some of segments considerably elongated, much longer than wide. Anal 
segment with limbate plate, spatulate, or funnellike (fig. 34, d, h, m). 
Prostomium hoodlike (fig. 34, 7, 1) or with limbate plate (fig. 34, b-c, g). 

Maldanidae (p. 302) 
Seaments not much longer than wide_..2 20. i225 2S 2ele lek lt 23 

23. Proboscis provided with dark, chitinous jaw pieces (fig. 31,7). Body smooth, 
elongate, cylindrical, resembling an earthworm. Parapodia weakly devel- 
Bped-cimple lobes <.s==-2-8) 000 LU Poms 282 Lumbrineridae (p. 275) 

Proboscis without chitinous jaws. Form variable____-_---------------- 24 

24. Body divided into 2 weakly to sharply separable regions, short anterior region, 
more or less flattened, with cushionlike neuropodia with several rows of 
setae, and long cylindrical posterior region with parapodial lobes projecting 


emt ye (hie ae,’ 6). Sa See 2) 2 eee Orbiniidae (p. 278) 
Parapodial lobes of median and posterior regions not projecting dorsally_. 25 
Deerostomium conical] ees. AvAe OTe 1D Se Ue ee Se eerie 4 26 
Paneponmurl otherwise 224-3 S524. 50726 Po _ OE Le SOU ou 27 

26. Body short, grub-shaped or lancet-shaped. Parapodia with simple capillary 
Permoraible: ithe ot) S84. a SA EW Opheliidae (p, 295) 


Body elongate, slender, fragile. Parapodia with capillary setae and, in long 
posterior region, with rows of hooded hooks on somewhat inflated tori 
Reena eet Ste. SU Le Ue eee oe Capitellidae (p. 298) 

27. Arborescent branchiae limited to anterior few segments. With capillary and 
forked: parapodial setae 2. aU S24 22-8225 tG_e Scalibregmidae (p. 293) 

Arborescent branchiae on long middle region of body. With capillary setae 

and sigmoid hooks embedded in elongated tori-_----- Arenicolidae (p. 300) 


Family POLYNOIDAE 


Prostomium bilobed, with two pairs of eyes in trapezoidal arrange- 
ment, three dorsal antennae, a pair of subulate ventral palps. First 
or tentacular segment with setae reduced in number (0-3), with two 
pairs of tentacular cirri (fig. 26, a). Parapodia biramous, with setae 
simple (not compound), paired dorsal scales or elytra on certain seg- 
ments and paired dorsal cirri on non-elytra-bearing segments, paired 
subulate ventral cirri (fig. 26, a, d). Pygidium or anal segment with 
pair of anal cirri. Paired segmental or nephridial papillae near 
ventral bases of parapodia. Muscular proboscis eversible, with circle 
of soft marginal papillae and two pairs of interlocking, chitinous jaws. 

Represented by 7 genera and 11 species. All the genera con- 
sidered here have 15 pairs of elytra on segments 2, 4, 5, 7, 9, 11, 13, 
15, 17, 19, 21, 23, 26, 29, 32; lateral antennae inserted ventral to 
median antenna on prostomium (fig. 26, a). 


914 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 108 


Key to the genera of Polynoidae from Point Barrow 


1. Notosetae stouter than or at least as stout as neurosetae, with tips blunt to 
pointed, not; eapillary. 2... 7.42. $8 231 sec estos ae 2 
Notosetae not as stout as neurosetae-___-___-._----------.-=----------- 5 

2. Notosetae large, smooth or with only faint transverse striations, few in number 
(5-9) netfee lS. WK on 5 ee eel ee te ee Melaenis (p. 214) 
Notosetae with transverse spinous rows, more numerous (more than 10)___ 3 

3. Neurosetae long, slender, at least some end in capillary tips_Antinoé (p. 215) 
Neurosetae stouter, with tips straight or slightly hooked, not capillary_____ 4 

4. All neurosetae with entire bare tips--._-.------------------ Eunoé (p. 216) 
Some of neurosetae bidentate, with secondary tooth well developed or rudi- 
montary (fig. | 265 %¢) 2ec)5 ers ae pope Sh ae a Harmothoé (p. 220) 

5. Some notosetae with capillary tips. Segments less than 40 (35-38), thus, 
posterior end of body covered with elytra____-.---------.------------ 6 
None of notosetae with capillary tips. Segments more than 40 (45-65), thus, 

a rather long posterior end without elytra. Elytra smooth, without 
tubercles=, 3 be .2hsccouieas select ee J ot ee Enipo (p. 225) 

6. Neurosetae of 2 kinds, upper few end in slender tips, rest end in bifid tips. 
Elytra smooth except for few microtubercles on anterior curved part. 
Arcteobia (p. 225) 

Neurosetae all similar, with entire tips. Elytra with microtubercles and 
sometimes also with macrotubercles_---..------------- Gattyana (p. 622) 


Genus Melaenis Malmgren, 1865 
Melaenis lovéni Malmgren, 1865 


Melaenis lovéni Malmgren, 1865, p. 78, pl. 10, fig. 10—Théel, 1879, p. 22.— 
Wirén, 1883, p. 391, pl. 28, fig. 4; pl. 29, fig. 3—Murdoch, 1885, p. 152.— 
Moore, 1908, p. 337.—Fauvel, 1914, p. 49.—Augener, 1928, p. 695.— 
Ditlevsen, 1937, p. 16.—Gorbunov, 1946, p. 38.—Wesenberg-Lund, 1950a, 
p. 9; 1950b, p. 29. 

Melaenis lovéni var. gigantea Wirén, 1883, p. 391, pl. 28, fig. 3; pl. 29, fig. 4.— 
Murdoch, 1885, p. 153. 

Description.—Length 25-76 mm., width including setae 9-22 mm- 
(up to 110 mm. long—Ditlevsen, 1937). Segments 39-41. Body 
elongated, thick, tapering both anteriorly and posteriorly. Pro- 
stomium without cephalic peaks, antennae smooth. Dorsal cirri 
with fine, scattered, clavate papillae. Elytra smooth, soft, trans- 
lucent to opaque, not covering middorsum and posterior few segments. 
Notosetae much stouter than neurosetae, few in number (5-9), 
light to dark amber-colored, smooth or with only very faint transverse 
striations. Neurosetae of two kinds: Numerous, long, slender, 
finely denticled, with capillary tips; few, shorter, with subequal, 
diverging, forked, blunt tips (one part of fork may be longer than the 
other). Cotor: In life and in alcohol: Wide greenish brown to 
bluish gray transverse stripes middorsally, green and white striped on 
dorsal tubercles, cirrophores and elytrophores; without color ventrally; 
elytra light to dark greenish in a wide, circular band, a circular area 


MARINE POLYCHAETE WORMS—PETTIBONE 215 


and lateral border without color; one specimen (collected October 4, 
1949) was noted as peach-colored from segment 10 posteriorly, the 
color showing mainly on the ventral side and extending into the 
parapodia (probably developing eggs). 

New records—Arcric AtasKA: Point Barrow base, washed ashore, 
1949 (July 30; August 21; September 22, 28, 30; October 4, 5, 16, 
17; 65 specimens); off Point Barrow base, along shore, 1.7 fms., on 
bottom of gravel with mud (2 specimens). East Coast Nortu 
America: Off Labrador, 45 fms., Blue Dolphin Expedition, 1949. 

Distribution —Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait, Greenland, Spitsbergen, Franz Josef 
Land, Kara Sea. Also Bering Sea; off Labrador. In 1.7-111 
fathoms. 


Genus Antinoé Kinberg, 1855 (sensu Malmgren, 1865) 


Antinoé sarsi Malmgren, 1865 


Antinoé sarst Kinberg, 1862 (MS.), p. 468 (fide Malmgren, 1865).—Malmgren, 
1865, p. 75, pl. 9, fig. 6 (part).—MclIntosh, 1900, p. 365, figs Chamberlin, 
1920, p. 8.—Hartman, 1944a, p. 334. 

Polynoé sarsi Théel, 1879, p. 16, pl. 1, fig. 8.—Wirén, 1883, p. 390.—Murdoch, 
1885, p. 152. 

Harmothoé sarsi Eliason, 1920, p. 20.—Annenkova, 1931, p. 203.—Friedrich, 
1939, p. 122.—Thorson, 1946, p. 48.—Wesenberg-Lund, 1950a, p. 7; 1950b, 
Dp: 2571901; p: 17. 

Harmothoé (Antinoélla) sarst Augener, 1928, p. 687. 

Antinoélla sarsi Annenkova, 1937, p. 153; 1938, p. 137.—Gorbunov, 1946, p. 38.— 
Zatsepin, 1948, p. 107, pl. 28, fig. 8. 


Description.—Length 21-68 mm., width including setae 11-27 mm. 
Segments 37 or 38. Fragments easily, posterior segments frequently 
regenerating. Body elongate, flattened, tapering slightly anteriorly 
and posteriorly. Prostomium with cephalic peaks distinct or poorly 
developed; anterior pair of eyes larger than posterior, slightly anterior 
to greatest width of prostomium. Antennae with short, clavate 
papillae. Dorsal cirri extending beyond setae, long, slender, tapering 
gradually, with short, clavate papillae. May bea single anal cirrus, 
extremely long, thick at base, tapering gradually; may be one long 
and one short anal cirrus. Elytra large, thin, soft, smooth, with 
scattered microtubercles and short, delicate, clavate papillae. Lose 
elytra readily; often show regenerating elytra, thus appearing variable 
in size. Notopodia and neuropodia extending into conspicuous, 
digitiform, acicular lobes. Setae yellowish or golden. Notosetae 
much thicker than neurosetae, large, clear basally; more distally, 
finely spinous, pointed. Neurosetae of two kinds, with capillary tips 
and with slender, relatively obtuse, slightly curved tips. Cotor: 
In life and preserved: Grayish green or light brown middorsally, 


216 PROCEEDINGS OF THE NATIONAL MUSEUM Vancate 
including elytrophores and dorsal tubercles; ventral surface without 
color or dusky midventrally; elytra gray, translucent to opaque, 
with a darker brownish-gray medial crescent-shaped area, with a 
colorless circular area medial to and a darker area posterior to the 
place of attachment. 

Remarks.—A. sarsi differs from A. badia (Théel), with which it has 
sometimes been confused, by having the neurosetae of two kinds, 
with capillary tips and with slender, relatively obtuse tips. In A. 
badia, all the neurosetae have capillary tips. 

Parasites —One of the 63 specimens had the parasitic copepod 
Herpyllobius arcticus Steenstrup and Liitken attached to the prosto- 
mium (identified by P. L. Illg). 

New records—Arctic ALASKA: Point Barrow base, washed ashore 
1949 (August 21, 24; September 1, 12, 20, 22, 24, 26; October 5, 16; 
58 specimens); off Point Barrow base, up to 5 miles from shore, 5 to 
30.7 fms., on bottoms of mud or stones, and in screen trap through 
hole in ice, 6 fms., April 11, 1949 (3 stations, 5 specimens); near 
Point Belcher, Icy Cape, off Point Barrow, Dall. Brrine Sra: 62°15’ 
N., 167°48’ W., 20 fms., G. M. Stoney, 1884. East Coast Nortu 
America: Off Labrador, 6-30 fms., Blue Dolphin Expeditions, 
1950, 1951. 

Distribution—Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Jan Mayen, Spitsbergen, Norway, Franz 
Josef Land, Barents Sea, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes to Great Britain, the Baltic; Labrador to Maine; Bering Sea; 
north Japan Sea. In 3-1,215 fathoms. 


Genus Eunoé Malmgren, 1865 


Key to the species of Eunoé from Point Barrow 


1. Elytra with scattered, bluntly conical microtubercles only, lacking fringe of 
papillae. Eyes small, anterior pair anterolateral on prostomium (scarcely 
visible, dorsally) 225 2 a2 3s ee ee ee ee KE. clarki 

Elytra with macrotubercles in addition to microtubercles, with lateral fringe 
of papillae. Eyes larger, anterior pair anterodorsal (visible dorsally)-_-- 2 

2. Body broad, oval, flattened. Extra rounded lobes on inner sides of elytro- 
phores and dorsal tubercles (lobes corresponding to elytrophores on non- 
elytra-bearing segments, fig. 26, c). Nuchal fold posterior to prostomium 
not prominent. Setae yellow. Elytra with microtubercles rather low, 
flattened, semiglobose, some bifid; macrotubercles confined essentially to 
single row near external border, nodular, with roughened tips. Antennae 
with’short papillae. [ie _ ere Se Coe en SOG TR E. nodosa 

Body more elongate, not so flattened. Without extra rounded lobes on inner 
sides of elytrophores and dorsal tubercles. Nuchal fold prominent. Setae 
dark amber-colored. Elytra with microtubercles one- to many-pronged; 
macrotubercles variable in number, size, and position, branched. Antennae 
with“ longer.“ paplllac ss 2eiee hee ae = es Se ee E. oerstedi 


MARINE POLYCHAETE WORMS—PETTIBONE 217 


Eunoé clarki Pettibone, 1951 
Eunoé clarki Pettibone, 1951, p. 44, fig. 1. 


Description—Length 36-38 mm., width including setae 12 mm. 
Segments 40 or 41. Body linear-oblong, tapered slightly anteriorly 
and posteriorly. Prostomium without cephalic peaks; anterior pair of 
eyes anterolateral. Antennae and dorsal cirri with short, clavate 
papillae. Elytra nearly cover dorsum, with conical microtubercles, 
without fringe of papillae. Conor: In alcohol: Middorsum trans- 
versely banded grayish green; elytra greenish gray, with a darker 
mottled pattern on most of the exposed parts, with a darker spot 
medial to a lighter area over the elytophore; dorsal cirri with pig- 
mented bands basally and subterminally. 

New record.—Arctic ALASKA: Point Barrow base, washed ashore, 
October 17, 1949 (2 specimens). 

Distribution.—Arctic Alaska. 


Eunoé nodosa (Sars, 1860) 
FicureE 26, ¢c 


Polynoé nodosa Sars, 1860, p. 58. 

Eunoé nodosa Malmgren, 1865, p. 64, pl. 8, fig. 4.—Fauvel, 1923, p. 51, fig. 18, 
a-e (part)—Annenkova, 1937, p. 149; 1938, p. 134.—Treadwell, 1937, p. 
27.—Friedrich, 1939, p. 122.—Berkeley and Berkeley, 1942, p. 187; 1943, 
p. 129.—Hartman, 1944a, pp. 334, 337—Gorbunov, 1946, p. 38.—Zatsepin, 
1948, p. 107, pl. 28, fig. 7. 

Eunoa nodosa Malmgren, 1867, p. 6.— Webster and Benedict, 1884, p. 700; 1887, 
p. 708.—MclIntosh, 1900, p. 291, figs. 

Polynoé scabra Théel, 1879, p. 7.—Wirén, 1883, p. 387, pl. 28, fig. 2; pl. 29, fig. 
1 (part; not pl. 28, fig. 1; not Aphrodita scabra Fabricius, 1780). 

Polynoé islandica Hansen, 1882, pp. 17, 24, pl. 1, figs. 15-21.—Murdoch, 1884, 
p. 152. 

Polynoé arctica Hansen, 1882, pp. 21, 27, pl. 3, figs. 1-5. 

Polynoé spinulosa Hansen, 1882, p. 28, pl. 1, figs. 6-10. 

Polynoé foraminifera Hansen, 1882, pp. 21, 23, 29, pl. 1, figs. 11-14. 

Harmothoé nodosa Moore, 1902, p. 271.—Ditlevsen, 1917, p. 6, pl. 3, fig. 10 (part; 
not pl. 2, fig. 1)—Augener, 1928, p. 684 (part).— Wesenberg-Lund, 1950a, 
p. 6 (part); 1950b, p. 17 (part); 1951, p. 10 (part). 

Eunoé depressa Hartman, 1948, p. 14 (part; not E. depressa Moore, 1905). 


Description —Length 13-75 mm., width including setae 7-35 mm. 
Segments 36 or 37. Prostomium with cephalic peaks short and blunt 
or lacking; anterior pair of eyes anterolateral, visible dorsally. An- 
tennae with short papillae. Dorsal cirri with long papillae. Body 
flattened ventrally, strongly arched dorsally, especially in anterior 
part. With characteristic bulbous structures on inner sides of bases 
of elytrophores and dorsal tubercles. Segmental papillae quite elon- 
gate, cylindrical, directed dorsally between the parapodia. Elytra 
with fringe of long papillae on external border. Elytral microtubercles 
numerous, close-set, rather low, flattened, semiglobose, some bifid; 


218 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


macrotubercles confined mostly to a single row near external border, 
dark colored to pale yellow, nodular, with roughened tips or a fasicle 
of short spikes. Elytra often covered with a good deal of debris and 
foreign material. Coztor: In life and in alcohol: Mediodorsal 
surface colorless or banded with olive-brown between elytrophores and 
dorsal tubercles; ventral surface without color or olive-brown; setae 
yellow or light amber-colored; elytra yellow or tannish mottled with 
reddish brown. 

Remarks.—Eunoé nodosa has been confused with E. oerstedi. The 
two species have been separated by Sars (1860), Malmgren (1865), 
Verrill (1881), Murdoch and Benedict (1885, as Polynoé islandica and 
P. scabra), and Treadwell (1937). They have been considered to be 
synonymous and a highly variable species by Théel (1879), Wirén 
(1883), Fauvel (1923), Ditlevsen (1917), Augener (1928), and Wesen- 
berg-Lund (1950a, b). Based on a study of the material from Point 
Barrow as well as numerous other specimens in the U. S. National 
Museum from Greenland and off New England, the two species appear 
to be separable on the basis of a number of characters as indicated 
in the key. They agree in having the prostomium with cephalic 
peaks short and blunt or lacking, the position of the eyes, the dorsal 
cirri with long papillae, and the palps each with six longitudinal rows 
of papillae. 

New records —Arctic ALASKA: Eluitkak Pass, Elson Lagoon near 
Point Barrow, stony-mud; off Point Barrow base, up to 12.1 miles 
from shore, 18.3-87 fathoms, on various combinations of mud, stones, 
pebbles, gravel, rocks (14 stations, 33 specimens). Brrine Sra: 57° 
N., 163° 48’ W., 38 fms., Alaska King Crab Expedition (Hartman 
(1948), as E. depressa); Albatross Sta. 3252, 57°22’ N., 164°24’ W., 
29.5 fms., black mud, 1890, and Sta. 3512, 57°49’ N., 169°27’ W., 
38 fms., 1893. SourHwxrsterRN ALASKA: Belkofsky Bay, 15-25 fms., 
Dall, 1880. Franz Joser Lanp: Aberdore Channel, east Alger 
Island, 10 fms., Baldwin-Ziegler Expedition, 1901. Easr Coasr 
Nortu America: Off Labrador, 10-95 fms., Blue Dolphin Expeditions, 
1949, 1950; Albatross Sta. 2432, off Newfoundland, 43°04’ N., 50°45’ 
W., 64 fms., 1885; Bay of Fundy, Grand Manan, Nova Scotia, Maine, 
Massachusetts, 16-120 fms., U. S. Fish Commission. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Jan 
Mayen, Spitsbergen, Norway, Franz Josef Land, Barents Sea, Novaya 
Zemlya, Kara Sea. Also Bering Sea; north Japan Sea; Iceland, Faroes, 
Shetlands to English Channel; Hudson Bay to Massachusetts. In 
10-690 fathoms. 


MARINE POLYCHAETE WORMS—PETTIBONE 219 


Eunoé oerstedi Malmgren, 1865 
FicureE 26, d 


Lepidonote scabra Oersted, 1843, p. 164, pl. 1, figs. 2, 7, 10, 12-13, 17-18 (not 
Aphrodita scabra Fabricius, 1780). 

Polynoé scabra Sars, 1860, p. 58.—Hansen, 1882, p. 11.—Wirén, 1883, p. 387, 
pl. 28, fig. 1 (part) Murdoch, 1885, p. 152 (not A. scabra Fabricius, 1780). 

Eunoé oerstedi Malmgren, 1865, p. 61, pl. 8, fig. 3—Moore, 1909b, p. 1385.— 
Sumner, 1913, p. 618.—Treadwell, 1937, p. 27.—Hartman, 1944a, pp. 334, 
337.—Gorbunov, 1946, p. 38.—Pettibone, 1953, p. 46, pl. 23. 

Eunoé barbata Moore, 1910, p. 334, pl. 28, figs. 1-6.—Treadwell, 1925, p. 1.— 
Hartman, 1939, p. 53.—Berkeley and Berkeley, 1945, p. 321; 1948, p. 14, 
fig. 16.—Pettibone, 1949, p. 2. 

Eunoé nodosa Fauvel, 1923, p. 51, fig. 18, a-e (part) Hartman and Reish, 1950, 
p. 7 (not Polynoé nodosa Sars, 1860). 

Harmothoé nodosa Ditlevsen, 1917, p. 6, pl. 2, fig. 1 (part; not pl. 3, fig. 10).— 
Augener, 1928, p. 684 (part) —Wesenberg-Lund, 1950a, p. 6 (part); 1950b, 
p. 17 (part); 1951, p. 10 (part; not P. nodosa Sars, 1860). 


Description.—Length 18-73 mm., width including setae 6-29 mm. 
Segments 37-39 (39-42 in specimens from Washington). Prostomium 
with cephalic peaks poorly developed or lacking; anterior pair of eyes 
anterolateral, visible dorsally. Antennae and dorsal cirri with long 
papillae. Body elongate, oval in cross section. Segmental papillae 
short and bulbous. Tips of notosetae may be worn down and blunt, 
especially in larger specimens. Elytra with lateral fringe of papillae. 
Elytral surface exceedingly scabrous, with microtubercles one- to 
many-pronged; macrotubercles branched, extremely variable in size, 
number, arrangement and shape; some brownish, some translucent. 
Cotor: In alcohol: Dusky, dark or greenish black between elytrophores 
and dorsal tubercles, with some color on cirrophores of dorsal cirri 
and tips of parapodia; setae dark amber-colored. 

Remarks.—This species is closely related to and has been confused 
with E. nodosa as discussed above. Examination of the type of £. 
barbata Moore from central California revealed no essential differences 
from EF. oerstedi. 

New records—Arctic Auaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 8 miles from 
shore, 18.3 to 75.5 fms., on bottoms of mud, sand, shells, pebbles, 
gravel, rocks, stones (10 stations, 20 specimens). CANADIAN ARCTIC: 
Baffin Island, 66°43’ N., 80°07’ W., 1927; center, south, and southeast 
corner Foxe Basin, 25-37 fms., 1927; Kneeland Bay in Frobisher 
Bay, Baffin Island, 7-12 fms., 1942; all collected by Bartlett. Wusr 
GREENLAND: Murchison Sound, 45-60 fms., Bartlett, 1938. East 
GREENLAND: Clavering Fiord, 1939; off Cape Hold with Hope, 
23-40 fms., Bartlett, 1939. Sourn Greentanp: Off Cape Farewell, 
70 fms., Bartlett, 1939. East Coast Norru America: Off Labrador, 


261112—54 


2 


220 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


25-40 fms., Blue Dolphin Expeditions, 1949, 1950; Nova Scotia, 
Grand Manan, Maine, Massachusetts, low water to 110 fms., U. S. 
Fish Commission. Japan: Albatross Sta. 3656, Hakodate Bay, 11.5 
fms., 1896. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Spits- 
bergen. Also from Norway to English Channel; Labrador to 
Massachusetts; Bering Sea to central California; and Japan. In 
low water to 516 fathoms. 


Genus Harmothoe Kinberg, 1857 


As used herein, Harmothoé includes Lagisca Malmgren, 1865; 
Evarne Malmgren, 1865; and Evarnella Chamberlin, 1919. 

Both species represented have the body short, attenuated posteriorly, 
segments about 40 (36-50), covered by elytra except for posterior 
few segments; elytra with numerous conical microtubercles. Pro- 
stomium with cephalic peaks prominent (fig. 26, a). Antennae and 
dorsal cirri with short papillae. Notosetae stouter than neurosetae, 
with long distal spinous regions and short, pointed, bare tips. 


Key to the species of Harmothoé from Point Barrow 


1. Anterior pair of eyes anteroventral on prostomium (slightly posterior and 
lateral to cephalic peaks), not visible dorsally (fig. 26, a). Elytra with 
or without large, globular macrotubercles near posterior border. 

H. imbricata 

Anterior pair of eyes anterolateral on prostomium, visible dorsally. Elytra 
with macrotubercles globular, sausage-shaped or elongate, rodlike. 

H. extenuata 


Harmothoe imbricata (Linné, 1767) 
FIGuRE 26, a, e 


Aphrodita imbricata Linné, 1767, p. 1804. 

Lepidonote cirrata Oersted, 1843, p. 166, pl. 1, figs. 1, 5-6, 11, 14-15. 

Polynoé cirrata Sars, 1860, p. 60. 

Harmothoé imbricata Malmgren, 1865, p. 66, pl. 9, fig. 8 Webster and Benedict, 
1884, p. 701; 1887, p. 709.—Johnson, 1897, p. 181; 1901, p. 390.—MclIntosh, 
1900, p. 314, figs—Moore, 1902, p. 270; 1903, p. 402; 1908, p. 334.—Dit- 
levsen, 1909, p. 7, pl. 2, fig. 5—Sumner, 1913, p. 617.—Southern, 1914, p. 
52.—Chamberlin, 1920, p. 4.—Eliason, 1920, p. 19.—Fauvel, 1923, p. 55, 
fig. 18, f-e; 1933, p. 10; 1936, p. 50.—Treadwell, 1925, p. 1; 1937, p. 26.— 
Augener, 1928, p. 677; 1939, p. 133.—Annenkova, 1934, p. 322; 1937, p. 151; 
1938, p. 186.—Okuda, 1938b, p. 83; 1939, p. 224.—Friedrich, 1939, p. 122.— 
Monro, 1939a, p. 345.—Hartman, 1944a, pp. 335, 337; 1948, p. 13.—Thorson, 
1946, p. 46, figs. 17-18.—Gorbunov, 1946, p. 38.—Zatsepin, 1948, p. 108, 
pl. 28, fig. 10.—Berkeley and Berkeley, 1948, p. 11, ‘fig. 9.—Pettibone, 1949, 
p. 2; 1953, p. 32, pls. 13-16.—Hartman and Reish, 1950, p. 6.—Wesenberg- 
Lund, 1950a, p. 6; 1950b, p. 18; 1951, p. 12. 


MARINE POLYCHAETE WORMS—PETTIBONE 921 


Polynoé imbricata Marenzeller, 1879, p. 9, pl. 2, fig. 1—Théel, 1879, p. 9—Hansen, 
1882, pp. 11, 13, 15, 23—Wirén, 1883, p. 389. 
Harmothoé levis Treadwell, 1937, p. 26, figs. 1-5. 

Description.—Length 24-53 mm., width including setae 8-19 mm. 
(length up to 65 mm.—Théel, 1879). Segments 37-39. Prostomium 
with anterior pair eyes ventral, slightly posterior and lateral to 
cephalic peaks. Elytra with scattered conical microtubercles, with 
or without few to numerous, small, brownish to reddish, globular to 
elongate cylindrical macrotubercles in one, two, or several irregular 
rows near posterior border; may be a few additional globular papillae 
in middle of elytra; with or without short elytral fringe of papillae. 
Neurosetae with long spinous regions, with long, bare, hooked tips, 
and usually with a subterminal tooth (may be broken off). Conor: 
In life and in alcohol: Irregularly pigmented middorsally, on cirro- 
phores, dorsal tubercles, and parapodial lobes; usually without color 
ventrally; elytra show remarkable color variations of mottled tan and 
brown, blackish to grayish, or reddish; inner halves of elytra red or 
grayish brown resulting in a middorsal, longitudinal, pigmented band; 
uniformly tan; tan with bilateral darker spots near places of attach- 
ment (the color variety of H. levis Treadwell). 

Remarks.—Examination of the types of H. levis Treadwell from 
Greenland waters revealed it to be but a color variation of H. 
imbricata. 

Parasites—Of the 215 specimens, 13 had parasitic copepods, 
Herpyllobius arcticus Steenstrup and Liitken, attached on the pro- 
stomium (identified by Paul L. Illg). 

New records—Arctic ALASKA: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms., gravel, stones, mud; Point Barrow base, 
washed ashore; off Point Barrow base, up to 16 miles from shore, 
11.7-123.5 fms., on bottoms of stones, mud, masses of worm tubes, 
and various combinations of mud, pebbles, gravel, stones, large 
perforated rocks, with barnacles, bryozoans, hydroids, tunicates, 
shells, worm tubes (34 stations, 215 specimens). CANADIAN ARTIC: 
Southeast corner Foxe Basin, 66°46’ N., 79°15’ W., 34-37 fms., 1927; 
shoal in Kneeland Bay, Frobisher Bay, Baffin Island, 7-12 fms., 
1942; west shore Frobisher Bay, 1942; off Daniels Island, northwest 
side in Newell Sound, Frobisher Bay, 10-30 fms., 1942; all collected 
by Bartlett. Wrsr Gremntanp: Oelricks Bay, mud, 1937; Vaigat, 
Disko Island, mud, 1937; between Capes Alexander and Chalon, 
25-40 fms., rocks, 1937; Walrus grounds, Murchison Sound, 45-60 
fms., 1938; off Conical Rock, 76° N., 67°30’ W., 20-40 fms., rocks, 
shell, 1938; all collected by Bartlett. Upernivik Harbor, 13 fms., 
U.S.S. Alert, 1884. Easr GreEnLAND: Clavering Fiord, 1939; 
Angmogssalik, 10-15 fms., 1939; off Cape Hold with Hope, 23-40 
fms., 1939; all collected by Bartlett. SprrsperGEn: Spitsbergen Sea, 


222 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


U.S.S. Alliance, 1881; South Gatt, northwest Spitsbergen, 79°40’ N., 
7 fms., E. Wilkinson. East Coast Norru America: Off Labrador 
and Newfoundland, intertidal to 60 fms., Blue Dolphin Expeditions, 
1949, 1950, 1951; off Nova Scotia, Grand Manan, Maine, Massachu- 
setts, Rhode Island, Long Island Sound, intertidal to 110 fms., U. S. 
Fish Commission. Brrine Sra: Bering Strait, 13 fms., Dall, 1880; 
62°54’ N., 166°38’ W., 22 fms., and 66°12’ N., 168°54’ W., Stoney, 
1884; St. George Island, Pribilofs, G. D. Hanna, 1914; Albatross 
Sta. 3252, 57°22’ N., 164°24’ W., 29.5 fms., 1890, and Sta. 3522, 
57°58’ N., 170°09’ W., 41 fms., 1893. SourHwestrern ALAsKa: Bay 
of Islands, Adak Island, 9-16 fms., 1873, and Coal Harbor, Unga 
Island, 1872, Dall. SourHrasterRN ALASKA: Wrangel, W. H. Jones. 
Sitka, “‘worm-eaten”’ wood, L. A. Beardslee. Japan: Albatross Sta. 
3656, Hakodate Bay, 1896. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Jan Mayen, Spits- 
bergen, Franz Josef Land, Barents Sea, Novaya Zemlya. Also 
Iceland and Norway to Mediterranean and Adriatic; Labrador to 
New Jersey; Bering Sea to southern California; and Japan. In low 
water to 2,030 fathoms. 


Harmothoé extenuata (Grube, 1840) 


Polynoé extenuata Grube, 1840, p. 86 (fide Claparéde, 1868).—Claparéde, 1868, 
p. 380, pl. 2, fig. 2. 

Polynoé rarispina Sars, 1860, p. 60.—Théel, 1879, p. 8—Hansen, 1882, p. 7.— 
Wirén, 1883, p. 389. 

Lagisca rarispina Malmgren, 1865, p. 65, pl. 8, fig. 2.—Verrill, 1881, pp. 311, 
314.—Webster and Benedict, 1884, p. 700; 1887, p. 709.—Moore, 1902, p. 
269; 1908, p. 335.—Fauvel, 1911, p. 9; 1914, p. 65.—Chamberlin, 1920, p. 
5.—Berkeley and Berkeley, 1943, p. 129; 1948, p. 16, figs. 18, 19. 

Lagisca propinqua Malmgren, 1867, p. 9, pl. 1, fig. 3, a-e. 

Lagisca extenuata Marenzeller, 1876, p. 133, pl. 1, fig. 1; 1904, p. 318.—McIntosh, 
1900, p. 307, figs.—Fauvel, 1923, p. 76, fig. 28, a-m; 1934a, p. 14.—Monro, 
1939a, p. 345.— Wesenberg-Lund, 1939a, p. 6, fig. 2; 1950a, p. 8; 1950b, p. 
27; 1951, p. 18. 

Lagisca impatiens Webster, 1879b, p. 102; 1886, p. 129, pl. 4, figs. 1-7. 

Polynoé semisculpta Hansen, 1882, p. 26, pl. 3, figs. 16-20 (?P. semisculptus 
Johnston, 1865). 

Lagisca floccosa McIntosh, 1900, p. 298, figs.—Southern, 1914, p. 51 (not Polynoé 
floccosa Savigny, 1820). 

Harmothoé extenuata Alaejos y Sanz, 1905, p. 55, pl. 9, fig. 8; pl. 10, figs. 1-12; 
pl. 11, fig. 1—Ehlers, 1913, p. 446.—Pettibone, 1953, p. 31. 

Harmothoé rarispina Ditlevsen, 1909, p. 5, pl. 1, figs. 2-4; 1937, p. 11—Augener, 
1928, p. 685.—Annenkova, 1937, p. 153; 1938, p. 1386.—Friedrich, 1939, p. 
122.—Gorbunov, 1946, p. 38.—Zatsepin, 1948, p. 108, pl. 28, fig. 15. 

Harmothoé triannulata Moore, 1910, p. 346, pl. 29, figs. 18-22.—Hartman, 1938a, 
p. 118.—Berkeley and Berkeley, 1948, p. 12, fig. 10. 

Lagisca extenuata var. spinulosa Fauvel, 1914, p. 64, pl. 4, figs. 27-29. 

Harmothoé propinqua Ditlevsen, 1917, p. 14, pl. 3, figs. 1, 3. 

Harmothoé rarispina forma propingua Annenkova, 1937, p. 153; 1938, p. 137. 


MARINE POLYCHAETE WORMS—PETTIBONE 223 


Evarnella triannulata Berkeley and Berkeley, 1942, p. 188; 1943, p. 130.—Hartman, 
1948, p. 13 (part; includes H. multisetosa Moore and H. extenuata).—Petti- 
bone, 1949, p. 1. 

Description.—Length 13-68 mm., width including setae 4-20 mm. 
(length up to 74 mm.—Ditlevsen, 1937). Segments 37-47. Prosto- 
mium with eyes large, anterior pair anterolateral, slightly anterior to 
widest part of prostomium, visible dorsally. Neurosetae with en- 
larged, long spinous regions, with tips slightly hooked, with small, 
secondary tooth present or absent; at least some of the setae show a 
remnant of a secondary tooth. The upper and lower neurosetae tend 
to be unidentate (not always) while the middle ones are bidentate, 
with a secondary tooth or remnant of it (in specimens from Woods 
Hole region only a few neurosetae have a secondary tooth or remnant 
of one). Elytra with numerous microtubercles, more or less uniform 
in size, conical, with tips blunt, pointed, or bifid; usually with few to 
fairly numerous macrotubercles distinctly set off from elytral surface, 
usually narrower at the base, brown, smooth, globular, sausage- 
shaped, or elongate fusiform, variable in number, 0-9 near posterior 
border, 0-13 scattered near center of elytron. Elytra with short 
fringe of papillae on external border. Cotor: In alcohol: Without 
color anteriorly; on posterior half, with brownish transverse bands and 
two brown spots basally on cirrophores (in specimens from Woods 
Hole region, found intertidally, the body is darkly pigmented grayish- 
green middorsally); antennae and dorsal cirri ringed with brown; 
elytra mottled with brown, sometimes with a darker spot posterior 
and medial to the place of attachment; the microtubercles may be 
covered with a brown extraneous material, giving astreaked appearance. 

Remarks.—H. extenuata is an extremely variable species as indicated 
by the number of names that have been applied to it. It reaches its 
greatest size in arctic waters; it is small when found intertidally, as 
in the Puget Sound region (up to 23 mm.—H. triannulata of Moore). 
When found intertidally off New England, it also is small (about 26 
mm.), and the secondary tooth of the neurosetae is suppressed, usually 
with only a remnant showing on a few neurosetae, the rest being entire 
(L. wmpatiens of Webster). In the collection from Point Barrow, 
where it was the most abundant polychaete, it is represented by 
specimens of all sizes from very small to large (up to 68 mm.). 

Parasites —Of the 3875 specimens, 3 had the parasitic copepod 
Herpyllobius arcticus Steenstrup and Lutken attached to the pros- 
tomium (identified by Illg). 

New records.—Arctic ALasKa: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 15 miles from 
shore, 13.3-123.5 fms., on bottoms of mud, stones, masses of worm 
tubes, and various combinations of mud, pebbles, rocks, gravel, 
stones, large perforated rocks, with shells, bryozoans, hydroids, 


224. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


worm tubes, from starfish and crab, Hyas coarctatus (37 stations, 375 
specimens). CanapIAN Arctic: Foxe Basin, 66°30’ N., 80° W., 
1927; center Foxe Basin, 25-31 fms., 1927; 7 miles east Cape Dor- 
chester, Foxe Channel, 25 fms., 1927; Baffin Island, 66°43’ N., 80°07’ 
W., 1927; Hurd Channel between Bushman Island and Melville 
Peninsula, 11 fms., rocky, 1933; south end Cape Martineau, Melville 
Peninsula, 7-15 fms., 1933; 3 miles south Salisbury Island, Hudson 
Strait, 27 fms., 1933; south end Cobourg Island, Baffin Bay, 75°40’ 
N., 78°58’ W., 11.3-20 fms., 1935; east end Cobourg Island, 75°40’ 
N., 78°50’ W., 23-36 fms., 1935; southeast corner Foxe Basin, 
66°46’ N., 79°15’ W., 34-37 fms., 1937; Kneeland Bay, Frobisher 
Bay, Baffin Island, 7-12 fms., 1942; all collected by Bartlett. Wausr 
GREENLAND: Vaigat, Disko Island, mud, 1937; between Capes 
Alexander and Chalon, 25-40 fms., rocky, 1937; Walrus grounds, 
Murchison Sound, 45 fms., 1938; all collected by Bartlett. Norra 
GREENLAND: North Omenolu near North Star Bay, 17 fms., and Cape 
Alexander, entrance to Smith Sound, rocky, 1932, Bartlett. Easr 
GREENLAND: Off Cape Hold with Hope, 23-40 fms., Bartlett, 1939. 
SoutH GREENLAND: Cape Farewell, 70 fms., Bartlett, 1939. Franz 
JosEF Lanp: Aberdore Channel east Alger Island, 10 fms., Baldwin- 
Ziegler Polar Expedition, 1901. Brrtne Sea: Alaska King Crab 
Investigation, 1941, Sta. D8-41, 58°34’ N., 165°17’ W., 42 fms.; 
Sta. D7-41, 57° N., 163°48’ W., 38 fms.; Sta. D11-41, 12 miles east 
Walrus Island, Pribilofs, 31-33 fms.; St. George Island, Pribilofs, 
village beach, 1914, and St. Paul Island, Pribilofs, 1915, Hanna; 
62°54’ N., 166°38’ W., 22 fms., and 62°15’ N., 167°48’ W., 20 fms., 
Stoney, 1884. Sourawestern AtasKa: Alaska King Crab Investiga- 
tion, 1940, 1941, Sta.: 89-40, Unga Strait, 37-47 fms.; 93-40, Spitz 
Island, 55-68 fms.; Sand Point; 128-40, Shelikof Strait off Hallo Bay, 
35-48 fms.; 60-40, Lenard Harbor, 20-25 fms.; L18-41, Kupreanof 
Strait, south side, 2 miles northwest Bare Island, 13-15 fms.; 72-40, 
Cold Bay, 15-50 fms.; 52-40, Canoe Bay, 35-40 fms., on Paralithodes 
camtschatica; 100-40, Alitak Bay, 30 fms.; 59-40, between inner 
Tliasik and Goloi Island, 20-30 fms. SourHrAsTERN ALASKA: 
Lituya Bay, 6-9 fms., Dall, 1874. Easr Coast Norra AMERICA: 
Off Labrador, 6-100 fms., Blue Dolphin Expeditions, 1949, 1950, 
1951; Bay of Fundy, Gulf of Maine, Massachusetts, Long Island 
Sound, 4-134 fms., U. S. Fish Commission; Woods Hole region, 
intertidal and dredged, Pettibone, 1950, 1951; Fish Hawk Sta. 8826, 
8898, Chesapeake Bay, 1920. 

Distribution—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Spits- 
bergen, Franz Josef Land, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Norway to Mediterranean and Adriatic; Hudson Bay to 
Chesapeake Bay; Bering Sea to southern California; north Japan Sea; 
South Africa. In low water to 1,000 fathoms. 


MARINE POLYCHAETE WORMS—PETTIBONE 235 


Genus Enipo Malmgren, 1865, sensu Levinsen, 1883 
Enipo gracilis Verrill, 1874 


Enipo gracilis Verrill, 1874a, pp. 407, 411, pl. 6, fig. 4; 1874b, pp. 347, 361, 378, 
pl. 5, fig. 3—Pettibone, 1953, p. 22, pl. 7. 

Enipo cirrata Treadwell, 1925, p. 1, figs. 1-4.—Hartman, 1938a, p. 120, fig. 39, 
d, e.—Berkeley and Berkeley, 1942, p. 188; 1945, p. 322; 1948, p. 16, fig. 20. 

Polynoé gracilis Hartman, 1942a, p. 26, figs. 23-26. 

Description.—Length 25-76 mm., width including setae 3-4 mm. 
Segments variable in number (45, with regenerating posterior end, 
to 74). Body elongate, slender, with sides nearly parallel. Prosto- 
mium without cephalic peaks. In some specimens from New England 
there are slight prostomial peaks (prostomial peaks present—Hart- 
man, 1942). Antennae and dorsal cirri with short, clavate papillae. 
Dorsal cirri short, barely reaching tips of setae in anterior part, longer 
in posterior region without elytra. Elytra oval, translucent, small, 
usually leaving middorsum uncovered, smooth, without papillae or 
tubercles. Notosetae form a short, bushy bundle, rather numerous 
(more than 30 in specimens from Point Barrow) to few (2-20 or 
rarely 0—Treadwell, 1925), slender to moderately stout, tapering 
gradually to short, bare, blunt tips, with spinous rows distally. 
Neurosetae stouter than notosetae, upper ones with longer spinous 
regions (17-23 or so rows), middle ones with shorter spinous regions 
(10 or so rows), with long, bare, slightly hooked tips (neurosetae may 
be all similar, with 5-10 transverse spinous rows). Cotor: In alcohol: 
Without color or brownish middorsally; elytra pigmented smoky 
brown on the medial halves. 

Remarks.—Known to be commensal with the maldanid Nicomache 
lumbricalis (Fabricius) in Alaska (as E. cirrata—Berkeley and Berke- 
ley, 1942) and off Halifax, Nova Scotia, and Cape Cod (specimens in 
the U. S. National Museum, identified by Verrill). 

New Records —Arctic Ataska: Off Point Barrow, up to 12.1 miles 
from shore, 21-123.5 fms., from breaking rock, masses of worm tubes, 
with stones, large perforated rocks (3 stations, 3 specimens). East 
Coast Norra America: Off Nova Scotia, Maine, Massachusetts, 2— 
198 fms., U. S. Fish Commission. 

Distribution —Alaskan Arctic; southwestern Alaska to Washington; 
Nova Scotia to Massachusetts. In 2—123.5 fathoms. 


Genus Arcteobia Annenkova, 1934 
Arcteobia anticostiensis (McIntosh, 1874) 


Eupolynoé anticostiensis McIntosh, 1874, p. 265, pl. 10, figs. 1-4.—Wirén, 1883, 
p. 390, pl. 29, fig. 2—Marenzeller, 1890, pl. 1. 
Harmothoé anticostiensis Augener, 1928, p. 691, pl. 11, fig. 13. 


226 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Arcteobia anticostiensis Annenkova, 1934, p. 322; 1937, p. 149, pl. 3, figs. 26, 27; 
1938, p. 133. 
Eucranta anticostiensis Hartman, 1944a, p. 337. 

Description.—Length 11-26 mm., width including setae 4-8 mm. 
Segments 35 or 36. Prostomium with distinct cephalic peaks; anterior 
pair of eyes anteroventral, not visible dorsally. Antennae and dorsal 
cirri with short, clavate papillae. Elytral pairs get larger posteriorly, 
the last pair usually extending beyond end of body. Elytra without 
fringe of papillae, smooth except for scattered microtubercles on 
anterior curved part. Upper notosetae shorter, stouter, with short, 
blunt to sharp-pointed tips; most of notosetae more slender, with 
capillary tips. Few upper neurosetae longer, ending in sharp, slender 
tips; most of neurosetae with bifid tips. Co tor: In life and in alcohol: 
Irregularly banded middorsally, greenish to greenish black, with a 
transverse stripe and often a pair of small spots; elytra with reddish 
brown pigmented area, usually C-shaped on inner and posterior parts 
and often a darker spot over place of attachment; may form a com- 
plete, circular, colored area. 

Remarks.—Probably commensal in habit; off Labrador, found com- 
mensal in the sinuous tubes of the terebellid Pista flexuosa (Grube), 
one worm per tube. 

New records —Arctic ALASKA: Point Barrow base, washed ashore; 
off Point Barrow base, up to 12.1 miles from shore, 24.7—123.5 fms., 
on bottoms of mud, masses of worm tubes, and various combinations 
of mud, gravel, stones, rocks, and large perforated rocks (7 stations, 
14 specimens). Easr Coast Norra America: Off Labrador, 40-95 
fms., Blue Dolphin Expedition, 1949; off Nova Scotia, Maine, Mas- 
sachusetts, 10-86 fms., U. S. Fish Commission. 

Distribution.—Alaskan Arctic; Labrador to Massachusetts; Bering 
Sea; north Japan Sea. In low water to 123.5 fathoms. 


Genus Gattyana McIntosh, 1897 
Key to the species of Gattyana from Point Barrow 


1. Notosetae with tips blunt to capillary. Anterior pair of eyes anteroventral, 
not visible dorsally; cephalic peaks distinct. Elytra with microtubercles 


All notosetae with capillary tips. Anterior pair of eyes anterolateral; cephalic 
peaks short and blunt. Elytra with conical and bifid microtubercles, with a 
few larger, knoblike macrotubercles.._ -....2_-.--se0_-2 eee G. ciliata 


Gattyana cirrosa (Pallas, 1766) 
FicureE 26, b 


Aphrodita cirrhosa Pallas, 1766, p. 95, pl. 8, figs. 3-6. 

Nychia cirrosa Malmgren, 1865, p. 58, pl. 8, fig. 1—Théel, 1879, p. 7.—Verrill, 
1881, pp. 306, 311.—Wirén, 1883, p. 387—Webster and Benedict, 1884, p. 
700; 1887, p. 708.—? Andrews, 1891, p. 279. 


MARINE POLYCHAETE WORMS—PETTIBONE 230. 


Gattyana cirrosa McIntosh, 1900, p. 285, figs——Moore, 1902, p. 259; 1908, p. 
337.—Southern, 1914, p. 51.—Eliason, 1920, p. 21.—Fauvel, 1923, p. 49, 
fig. 17, a-g.—Augener, 1928, p. 692; 1939, p. 136.—Gustafson, 1936, p. 5.— 
Treadwell, 1937, p. 25——Annenkova, 1937, p. 148; 1938, pp. 83, 96, 133, 
222.— Monro, 1939a, p. 345.—Berkeley and Berkeley, 1943, p. 129; 1948, 
p. 13, fig. 13—Thorson, 1946, p. 45—Gorbunov, 1946, p. 38.—Hartman, 
1948, p. 14.—Zatsepin, 1948, p. 106, pl. 28, fig. 5——Pettibone, 1949, p. 2; 
1953, p. 41, pl. 20.—Wesenberg-Lund, 1950a, p. 5, pl. 1, figs. 1, 2; 1950b, 
p. 12; 1951, p. 9. 

Gattyana cirrhosa Chamberlin, 1920, p. 8. 

Description.—Length 11-35 mm., width including setae 4-12 mm- 
Segments 35-38. Elytra with long papillae on posterior and external 
borders as well as scattered over the exposed part of elytral surface, 
usually covered with debris, giving a straggly appearance. Elytral 
tubercles amber-colored, exceedingly variable in shape, conical or 
cylindrical, simple, bifid or quadrifid, showing various degrees of 
being worn down; tubercles on the anterior few pairs of elytra tend 
to be less worn down, showing their more characteristic shapes. 
Cotor: In alcohol: Elytra tan mottled with brown or uniformly tan; 
often with blackish foreign material on elytral tubercles and papillae. 
Specimens washed ashore were essentially free from debris, the elytra 
whitish, iridescent, with amber-colored tubercles and inconspicuous, 
delicate papillae. 

Parasites—Three of the 126 specimens had the parasitic copepod 
Herpyllobius arcticus Steenstrup and Liitken attached to the prostomia 
(identified by Illg). 

New records—Arctic Ataska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; Point Barrow base, washed ashore; off Point 
Barrow base, up to 15 miles from shore, 18.3-123.5 fms., on bottoms 
of mud, stones, masses of worm tubes, and various combinations of 
mud, pebbles, small and large rocks, stones, gravel, and worm tubes; 
from breaking rock (30 stations, 126 specimens). Cape Smyth, 2.5-3 
fms., Point Barrow Expedition, 1883. Canapian Arctic: South end 
Cape Martineau, Melville Peninsula, 7-15 fms., 1933; Foxe Basin, 
1933; east end Cobourg Island, Baffin Bay, 75°40’ N., 78°50’ W., 
23-40 fms., 1935; Kneeland Bay, Frobisher Bay, Baffin Island, 14 
fms., 1942; all collected by Bartlett. Wurst Grerntanp: Vaigat, 
Disko Island, 1937; between Capes Alexander and Chalon, 25-40 fms., 
rocky, 1937; Walrus grounds, Murchison Sound, 60 fms., 1938; 
northwest Conical Rock, 1938; all collected by Bartlett. Upernivik 
Harbor, 13 fms., and Godhavn, U. S. S. Alert, 1884. East GrreEn- 
LAND: Off Cape Hold with Hope, 4-6 fms., Bartlett, 1939. Norra 
GREENLAND: North Omenolu near North Star Bay, 17 fms., Bartlett, 
1932. SprrsBERGEN: Spitsbergen Sea, U.S. S. Alliance, 1881. Brrine 
Sra: Bering Strait, 13 fms., Dall, 1880. Albatross Sta. 3326, 53°40’ 


228 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


N., 167°41’ W., 1890. SourHwestern AtasKa: Iluiliuk Harbor, 
Unalaska, Dall, 1871. SouruHmasteRN ALASKA: Sitka, Dall. Easr 
Coast Norru America: Off Labrador, 13-125 fms., Blue Dolphin 
Expeditions, 1949, 1950, 1951. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Novaya 
Zemlya. Also Iceland, Faroes, Norway to France; Hudson Bay to 
Massachusetts; Bering Sea to Washington; north Japan Sea. In 
low water to 630 fathoms. 


Gattyana ciliata Moore, 1902 


Gatiyana ciliata Moore, 1902, p. 263, pl. 13, figs. 14-19; pl. 14, fig. 20; 1905a, 
p. 525; 1908, p. 337.—Annenkova, 1937, p. 148, pl. 2, figs. 9, 11; pl. 4, fig. 33; 
1938, pp. 83, 100, 132, 224.—Berkeley and Berkeley, 1948, p. 12, fig. 11.— 
Pettibone, 1953, p. 40, pl. 19. 

Description.—Length 63-65 mm., width including setae 25-29 mm. 
(length up to 80 mm.—Annenkova, 1937). Segments 36 or 37. 
Cephalic peaks lacking or short and blunt. Elytral tubercles of 
several kinds: Microtubercles numerous, conical, pointed, hooked, 
some bifid; usually intermediate-sized tubercles elongated, conical, 
some bifid, in several rows arranged diagonally from center of elytron 
laterally; conical macretubercles with blunt, roughened tips, usually 
near posterior border of elytra, variable in number, size, and shape. 
Conor: In alcohol: Elytra tannish mottled with brown. 

New records —Anrctic ALasKa: Off Point Barrow base, up to 12.1 
miles from shore, 20-87 fms., on bottoms of rocks, stones, small 
amount of gravel (4 stations, 4 specimens). Brrtne Sua: Albatross 
Sta. 3606, 55°27’ N., 167°47’ W., 87 fms., green mud, fine sand, 1895. 

Distribution.—Alaskan Arctic (originally recorded from Greenland 
waters, corrected to Icy Cape, Alaska—Moore, 1902, 1905); Bering 
Sea to Washington; north Japan Sea. In 4.4-303 fathoms. 


Family SIGALIONIDAE 


Prostomium subglobular, with two pairs of eyes, dorsal antennae 
1-3, a pair of subulate, ventral palps (fig. 26, f). First or tentacular 
segment with or without numerous setae (lacking in Pholoé), with 
two pairs of tentacular cirri. Parapodia biramous, with notosetae 
simple, neurosetae simple or composite or both; with paired dorsal 
elytra on certain segments; without dorsal cirri; with paired subulate 
ventral cirri. Pygidium with pair of anal cirri. With or without 
cirriform branchiae and ciliated cushions or ctenidia on parapodia 
(both lacking in Pholoé). Muscular proboscis eversible, with circle of 
soft marginal papillae and two pairs of interlocking, chitinous jaws. 

Represented by a single species from Point Barrow. 


MARINE POLYCHAETE WORMS—PETTIBONE 229 


= 
rawr 


né---------- 


meee 


Ficure 26.—Polynoidae: a, Harmothoé imbricata, dorsal view anterior end, first pair elytra 
removed; b, Gattyana cirrosa, part of elytron enlarged, showing elytral tubercles and 
papillae; c, Eunoé nodosa, dorsal view part of two segments showing extra rounded lobes 
on inner sides of elytrophore and dorsal tubercle; d, Eunoé oerstedi, biramous parapodium; 
e, Harmothoé imbricata, bidentate neuroseta. Sigalionidae: f, Pholoé minuta, dorsal 
view anterior end, first pair elytra removed. (For explanation of symbols, see p. 210.) 


230 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Genus Pholoé Johnston, 1839 
Pholoé minuta (Fabricius, 1780) 


FIcure 26, f 


Aphrodita minuta Fabricius, 1780, p. 314. 

Aphrodita longa Fabricius, 1780, p. 313. 

Pholoé minuta Oersted, 1848, p. 169, pl. 1, figs. 3, 4, 8, 9, 16—Malmgren, 1865, 
p. 89, pl. 11, fig. 13.—Théel, 1879, p. 24.—Webster and Benedict, 1884, 
p. 701; 1887, p. 709.—MclIntosh, 1900, p. 437, figs——Moore, 1902, p. 274; 
1908, p. 338.—Ehlers, 1913, p. 450.—Southern, 1914, p. 57—Chamberlin, 
1920, p. 5.—Eliason, 1920, p. 22.—Fauvel, 1923, p. 120, fig. 44, a-h.— 
Augener, 1928, p. 673.—Annenkova, 1934, p. 322; 1937, p. 155; 1938, p. 138.— 
Berkeley and Berkeley, 1942, p. 189.—Hartman, 1944a, pp. 335, 337; 1948, 
p. 15.—Thorson, 1946, p. 49, fig. 19; Gorbunov, 1946, p. 38.—Zatsepin, 
1948, p. 109, pl. 28, fig. 17—Hartman and Reish, 1950, p. 8.—Wesenberg- 
Lund, 1950a, p. 9; 1950b, p. 31; 1951, p. 22.—Pettibone, 1953, p. 77, pl. 39. 

Pholoé tuberculata Southern, 1914, p. 57, pl. 6, fig. 14, A-L.—Berkeley and 
Berkeley, 1945, p. 323; 1948, p. 22, fig. 26. 


Description—Length 5-25 mm., width including setae 1-4 mm. 
Segments 36-84. Body small, elongate, nearly linear, flattened 
dorsoventrally; fragments easily. Lateral ventral surface and ventral 
sides of parapodial lobes thickly set with small, globular to elongate 
papillae, usually covered with debris. Prostomium small, oval; 
anterior and posterior pairs of eyes closely approximated. Single 
median antenna short, subulate (lateral antennae lacking). Ten- 
tacular segment achaetous; tentacular cirri similar to median antenna. 
A digitiform facial tubercle on a rounded lobe dorsal to the mouth 
and ventral to the prostomium. (On this character, Southern (1914) 
differentiated P. tuberculata; the lobe, on which the facial tubercle is 
located, appears to be retractile; surrounded by the bases of the palps, 
tentacular cirri and median antenna, the facial tubercle is easily 
overlooked unless the region is quite relaxed.) Elytra numerous, on 
segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 24, continuing on all 
segments to end of body; elytra nearly cover the dorsum except for a 
narrow middorsal part, usually pressed close to body; elytral surface 
smooth, with few microtubercles, somewhat areolate as seen under 
magnification, posterior border with few scattered papillae, somewhat 
moniliform. Notosetae simple, slender, finely spinous, tapering to 
capillary tips, upper ones shorter, with pronounced angles, the rest 
longer, with more gradual curves. Neurosetae stouter than notosetae, 
compound, with short, falcate terminal pieces. Conor: In alcohol: 
Without color or irregularly pigmented greenish gray; elytra trans- 
lucent, colorless, or greenish gray with a light spot over the place of 
attachment, or brownish; often with much debris. 


MARINE POLYCHAETE WORMS—PETTIBONE 931 


New records—Arctic AtasKa: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Pomt Barrow base, up to 12.1 miles from 
shore, 16.7—123.5 fms., on bottoms of stones, masses of worm tubes, 
and various combinations of mud, gravel, stones, rocks, large per- 
forated rocks, with bryozoans, hydroids; from screen trap through 
hole in ice, mud (16 stations, 30 specimens). Wrst GREENLAND: 
Vaigat, Disko Island, mud, 1937, and Walrus grounds, Murchison 
Sound, 45 fms., 1938, Bartlett. Easr Coast Nortu America: Off 
Labrador, 6 fms., Blue Dolphin Expedition, 1949; off Nova Scotia, 
St. Georges Bank, Maine, Massachusetts, Rhode Island, intertidal to 
110 fms., U. S. Fish Commission. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Jan Mayen, Norwegian 
Sea, Spitsbergen, Franz Josef Land, Novaya Zemlya, Kara Sea. 
Also Iceland, Faroes, Shetlands, Norway to northern France; Labrador 
to Rhode Island; Bering Sea to southern Oregon; north Japan Sea; 
off South Africa. In low water to 1,254 fathoms. 


Family PaHyLLopocIDAE 


Prostomium subconical, suboval or cordiform, with two eyes, four 
or five antennae, without palps (fig. 27, a-d). Anterior segments 
1-3 modified, with two to four pairs tentacular cirri. Parapodia 
uniramous (exceptionally biramous); setae compound (some may be 
simple). Dorsal and ventral cirri foliaceous or more or less globular 
(fig. 27, e~). Two anal cirri. Proboscis eversible, with papillae, 
unarmed. Active; mucus secreted in quantities. 

Represented by four genera and seven species. All genera repre- 
sented have uniramous parapodia, with compound setae. 


Key to the genera of Phyllodocidae from Point Barrow 


1. Tentacular cirri 4 pairs; first tentacular segment rudimentary, not visible 
dorsally, with 1 pair tentacular cirri lateral to prostomium; second segment 
distinct, with 2 pairs tentacular cirri; third segment distinct, with 1 pair 
tentacular cirri, 1 pair normal ventral cirri (fig. 27, c, d)__.-..__.______-- 2 

Tentacular cirri 3 pairs; first tentacular segment distinct dorsally, with 1 pair 
tentacular cirri; second segment distinct, with 2 pairs tentacular cirri (fig. 
27, b). Prostomium oval, without occipital tubercle_____ Mystides (p. 232) 

Tentacular cirri 2 pairs on first achaetous segment (fig. 27, a). Without dorsal 
cirri on second segment which has a setigerous lobe (may be very small or 
lacking) and a foliaceous ventral cirrus. Prostomium somewhat triangular, 
widest basally, with anterior part rounded, with 2 pairs short, subulate, 
frontal antennae, usually with 2 deep-set eyes (may not be visible when 
preserved), usually with a median occipital depression containing a small 
occipital tubercle (not always conspicuous), with a pair of lateral nuchal 
Er er a VERS OREM” Oa tae Oe OP PERE faye Cee peat 5)e ee eee Eteone (p. 232) 


232 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


2. Antennae 4 (2 pairs frontal antennae). Prostomium cordiform, with an 
occipital tubercle in the posterior notch (fig. 27, d). Proboscis with papillae 
proximally arranged in longitudinal rows__----------- Phyllodoce (p. 236) 

Antennae 5 (2 pairs frontal antennae and unpaired median antenna). Prosto- 
mium oval or bluntly conical, without occipital tubercle (fig. 27, ec). Pro- 
boscis with surface smooth, wrinkled, or with few scattered papillae 

Eumida (p. 238) 


Genus Mystides Théel, 1879 
Mystides borealis Théel, 1879 


Fiacure# 27, 6 


Mystides borealis Théel, 1879, p. 35, pl. 2, figs. 29-32.—Southern, 1914, p. 72, 
pl. 8, fig. 19, a-p.—Bergstrém, 1914, p. 176, fig. 64.—Fauvel, 1923, p. 181, 
fig. 65, a-d.—Augener, 1928, p. 711.—Zatsepin, 1948, p. 113, pl. 29, fig. 15.— 
Wesenberg-Lund, 1950b, p. 37. 

Mystides viridis Webster and Benedict, 1887, p. 712, pl. 1, figs. 10, 11, 13; pl. 2, 
fig. 12. (Type in USNM.) 

Mystides notialis Ehlers, 1913, p. 457, pl. 29, figs. 1-4. 

Description.—Length 5-9 mm., width 0.6 mm. Segments 33-64 
(up to 16 mm. long and 73 segments—Ehlers, 1913). Body small, 
linear, tapering anteriorly and posteriorly, flattened dorsoventrally. 
Prostomium oval], wider than long, with two deeply-set eyes in middle 
of prostomium, with four long, filiform antennae about one-third 
way back from anterior tip. Tentacular cirri 3 pairs, enlarged basally, 
with long, filiform tips. Dorsal, ventral, and anal cirri oval, thick, 
flattened. Cotor: In alcohol: Without color or irregularly pigmented 
with brown; cirri deep reddish brown. 

Remarks.—The description of M. notialis from the Antarctic by 
Ehlers (1913) follows closely that of MM. borealis and is herein considered 
to be synonymous. 

New records —Arctic AuAsKa: Off Point Barrow, 7.5 miles from 
shore, 36 fms., on bottom of stones, perforated rocks (1 station, 2 
specimens). West Coast Nort America: Off Flat Point, Lopez 
Island, Washington Sound, Pettibone (1 specimen). 

Distribution.—Scattered records in the Arctic: Arctic Alaska, West 
Greenland, Spitsbergen, Novaya Zemlya. Also Ireland, Madeira, 
Mediterranean; east coast North America (Maine); west coast North 
America (Washington Sound); Antarctic (Kerguelen). In 4-214 
fathoms. 


Genus Eteone Savigny, 1817 


The four species represented have the body elongate, somewhat 
flattened dorsoventrally, tapering anteriorly and posteriorly, with 
segments numerous (100-300). Proboscis with a circle of soft papillae 
around opening. 


MARINE POLYCHAETE WORMS—PETTIBONE Dee 


Key to the species of Eteone from Point Barrow 


1. Proboscis with 2 lateral, longitudinal rows of numerous soft papillae. Dorsal 
pair tentacular cirri slightly longer or up to 2-3 times longer than ventral 
pair. Dorsal cirri wider than long, nearly symmetrical, subcircular or 
slightly lanceolate (fig. 27, e). Color in alcohol: Darkly pigmented with 
bluish-violet iridescence, with longitudinal, lighter colored bands on each 
side of median dorsal line (or 3 dorsal longitudinal dark bands—1 median 
and 2 lateral). Anal cirri subcylindrical, 3—4 times longer than wide. 
Second segment with setigerous lobe and setae well developed__._E. barbata 

Surface of proboscis smooth, rugose or irregularly papillate. Two pairs ten- 
tacular cirri subequal or ventral pair somewhat longer than dorsal pair___ 2 

2. Dorsal cirri small, not much larger than parapodial lobes, longer than or as 
long as wide, almost symmetrical, thick, flattened, bluntly conical (fig. 27, 
h). Anal cirri short, thick, almost spherical. Second segment with seti- 
gerous lobe and setae well developed__-_-_--------------------- E. longa 

Dorsal cirri much larger than parapodial lobes, wider than long__--------- 3 

3. Dorsal cirri slightly asymmetrical, oval (fig. 27, g). Color in alcohol: Dorsum 
brownish with bluish-violet irridescence. Anal cirri short, thick, almost 
spherical. Second segment with setigerous lobe and setae well developed. 

E. flava 

Dorsal cirri much wider than long (about 2.5 times), asymmetrical, obliquely 

oval, with lower part larger than upper part (fig. 27, f). Color in alcohol: 

Middle third of dorsum light brownish, lateral third deep reddish brown. 

Anal cirri elongate, cylindrical, 3-4 times longer than wide. Second segment 
with setigerous lobe and setae poorly developed or lacking. 

E. spetsbergensis 


Subgenus Mysta Malmgren, 1865 
Eteone (Mysta) barbata (Malmgren, 1865) 
FIGURE 27, e 


Mysta barbata Malmgren, 1865, p. 101, pl. 15, fig. 34; 1867, p. 26, pl. 3, fig. 20.— 
Bergstrém, 1914, p. 207, fig. 79—Southern, 1914, p. 75.—Eliason, 1920, 
p. 28.—Augener, 1928, p. 711; 1939, p. 186.—Zatsepin, 1948, p. 113, pl. 29, 
fig. 14—Wesenberg-Lund, 1951, p. 31. 

Eteone striata Ditlevsen, 1917, p. 66, pl. 5, figs. 11, 17, 19. 

Eteone (Mysta) barbata Fauvel, 1923, p. 176—Annenkova, 1937, p. 159; 1938, 
p. 146.—Berkeley and Berkeley, 1942, p. 190.—Thorson, 1946, p. 62, fig. 
27, A-B. 

Description.—Length 42-180 mm.; width without setae 2.5-7 mm. 
See key. Cotor: In life: Three iridescent, brownish-purple dorsal 
longitudinal bands, median and lateral. 

New records —Arctic ALASKA: Eluitkak Pass, Elson Lagoon near 
Point Barrow, stony-mud; off Point Barrow base, 6.1-27 fms., up 
to 3.2 miles from shore, on bottoms of mud, gravel, stones, rocks; 
also caught in trap on bottom (3 stations, 4 specimens). 


934 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Distribution.—Scattered records in the Arctic: Siberian and Alaskan 
Arctic, Spitsbergen, Novaya Zemlya, Kara Sea. Also Iceland, Nor- 
way to Ireland, North Sea, Baltic; Bering Sea; north Japan Sea. 
In 5.5-60 fathoms. 


Eteone longa (Fabricius, 1780) 
FIGURE 27, h 


Nereis longa Fabricius, 1780, p. 300. 

Eteone longa Oersted, 1843, p. 185, pl. 2, figs. 20, 28 —Bergstrém, 1914, p. 192, 
fig. 72.—Chamberlin, 1920, p. 11.—Eliason, 1920, p. 26.—Fauvel, 1923, 
p. 172, fig. 62, a-d; 1933, p. 16—Augener, 1928, p. 710; 1939, p. 136.— 
Annenkova, 1934, p. 322; 1937, p. 158; 1938, p. 145.—Thorson, 1946, p. 
59, fig. 26, a-z.—Gorbunov, 1946, p. 38.—Zatsepin, 1948, p. 113, pl. 29, 
fig. 13.—-Berkeley and Berkeley, 1948, p. 41, figs. 57, 58 — Wesenberg-Lund, 
1950a, p. 11, pl. 2, figs. 6, 7; 1950b, p. 38; 1951, p. 29. 

Eteone arctica Malmgren, 1867, p. 27, pl. 2, fig. 12. 

Eteone cinerea Webster and Benedict, 1884, p. 705, pl. 1, figs. 1-5. 

Eteone tuberculata Treadwell, 1922, p. 174, figs. 7-10. 

Eteone californica Hartman, 1936a, p. 131, figs. 43-46; 1948, p. 20, fig. 4, a-d.— 
Rioja, 1941, p. 687. 

Description.—Length 10-65 mm., width without setae 0.5-3 mm. 
(length up to 160 mm.—Berkeley and Berkeley, 1948). See key. 
Cotor: In life: White. 

Remarks.—E. tuberculata Treadwell (1922) from Friday Harbor, 
Washington, FE. californica Hartman (1936) from central California 
(type examined as well as specimens from southeastern Alaska— 
Hartman, 1948), and E. cinerea Webster and Benedict (1884) from 
Provincetown, Massachusetts (type in USNM), appear to be 
synonymous with LE. longa. 

New records.—Arctic AuasKA: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, 1.7—75.5 fms., up to 8 
miles from shore, on bottoms of sandy-mud, stones, and various 
combinations of mud, pebbles, stones, gravel, rocks (12 stations, 40 
specimens). Wersr Coast Nort America: San Juan Archipelago, 
Washington, Pettibone. Wexst GreEen.aAnp: Vaigat, Disko Island, 
mud, Bartlett, 1937. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Novaya Zemlya. Also Iceland, Norway to English Channel; 
Hudson Bay to Massachusetts; Bering Sea to México; north Japan 
Sea; China. In low water to 518 fathoms. 


MARINE POLYCHAETE WORMS—PETTIBONE 235 


Eteone flava (Fabricius, 1780) 
FIGURE 27, g 


Nereis flava Fabricius, 1780, p. 299. 

Eteone flava Malmgren, 1865, p. 102, pl. 15, fig. 35—Bergstrém, 1914, p. 196, 
fig. 74.—Eliason, 1920, p. 27.—Fauvel, 1923, p. 173, fig. 62, e, f—Augener, 
1928, p. 709.—Annenkova, 1937, p. 158; 1938, p. 145.—Zatsepin, 1948, p. 
113, pl. 29, fig. 12—Wesenberg-Lund, 1950a, p. 11, pl. 2, fig. 7, ¢; 1950b, p. 
39; 1951, p. 30. 

Eteone depressa Malmgren, 1865, p. 103, pl. 15, fig. 36.—Southern, 1914, p. 79. 

Eteone sarsi Malmgren, 1867, p. 28, pl. 2, fig. 14.—Webster and Benedict, 1887, 


A 
sins lentigera Malmgren, 1867, p. 29, pl. 2, fig. 13. 

Description —Length 24-68 mm.; width without setae 1-4 mm. 
(length up to 120 mm.—Fauvel, 1923). See key. Conor: In life: 
Pinkish flesh and grayish white. 

New records:—Arctic ALASKA: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, 8.3-54.6 fms., up to 
7.5 miles from shore, on bottoms of mud, and various combinations 
of mud, stones, gravel, rock, large perforated rocks, shells (10 stations, 
22 specimens). 

Distribution.—Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen, Novaya Zemlya. Also 
Iceland, Faroes, Norway to English Channel; east coast North Amer- 
ica (Maine); Bering Sea; north Japan Sea. In low water to 471 
fathoms. 

Eteone spetsbergensis Malmgren, 1865 


Figure 27, f 


Eteone spetsbergensis Malmgren, 1865, p. 102, pl. 15, fig. 38.—Théel, 1879, p. 31, 
pl. 2, figs. 21, 22.—Bergstrém, 1914, p. 202, fig. 77.—Ditlevsen, 1917, p. 66, 
pl. 5, figs. 12, 14, 18 —Eliason, 1920, p. 27.—Augener, 1928, p. 708.—Annen- 
kova, 1937, p. 159; 1938, p. 145.—Hartman, 1948, p. 20, fig. 5, b— Berkeley 
and Berkeley, 1948, p. 42.—Zatsepin, 1948, p. 113, pl. 29, fig. 11— Wesen- 
berg-Lund, 1951, p. 31. 

Description.—Length 17-75 mm., width without setae 0.7-4 mm. 
(length up to 100 mm.—Augener, 1928). See key. Conor: In life: 
Pale green ventrally, reddish brown dorsolateral bands, with dark 
dorsal cirri; orange colored eggs. 

New records—Anrctic ALASKA: Off Browerville near Point Barrow; 
off Point Barrow base, 1.7-36 fms., up to 4% miles from shore, on 
bottoms of sandy-mud and various combinations of mud, gravel, 
rocks, stones, shell (4 stations, 4 specimens). 

Distribution.—Scattered records in the Arctic: Arctic Alaska, Spits- 
bergen, Franz Josef Land, Novaya Zemlya. Also Iceland, Norway 
to Scotland; Bering Sea; western Canada; north Japan Sea. In 1.7- 
40 fathoms. 


261112—54——_3 


236 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Eteone sp. larvae 


Larvae of a species of Eteone found in a mass of jelly (off Point 
Barrow base, 1.7 fms., 300 ft. out, on sandy-mud, July 20, 1948); 11 
segments, of which 8 were setigerous; triangular prostomium with 
short antennae and eyes; tentacular segment with short tentacular 
cirri; anal cirri short, spherical. 


Genus Phyilodoce Savigny, 1817 
Subgenus Anaitides Czerniavsky, 1882 


Phyllodoce (Anaitides) groenlandica Oersted, 1842 


Figure 27, d, i 


Phyllodoce groenlandica Oersted, 1842, p. 121 (fide Bergstrém, 1914); 1848, p. 192, 
pl. 2, figs. 19, 21, 22, 29-32.— Malmgren, 1865, p. 96; 1867, p. 21, pl. 2, fig. 
9.—Webster and Benedict, 1884, p. 703; 1887, p. 710.—Murdoch, 1885, p. 
153.—Moore, 1903, p. 428.—MclIntosh, 1908, p. 86, figs—Ditlevsen, 1917, 
p. 56.—Fauvel, 1923, p. 153, fig. 54, f-i—-Augener, 1928, p. 703.—not 
Treadwell, 1937, p. 28 (=Paranaitis wahlbergi (Malmgren), in USNM).— 
Annenkova, 1937, p. 156.—Friedrich, 1939, p. 122.—Thorson, 1946, p. 52, 
fig. 21.—Gorbunov, 1946, p. 38.—Berkeley and Berkeley, 1948, p. 46, fig. 
66.—Wesenberg-Lund, 1950a, p. 10; 1950b, p. 32; 1951, p. 26. 

Phyllodoce citrina Moore, 1908, p. 328.—Hartman, 1948, p. 19 (part; includes 
P. groenlandica and P. mucosa, in USNM; not P. citrina Malmgren, 1865). 

Anaitides groenlandica Bergstrém, 1914, p. 141, fig. 42—Southern, 1914, p. 68.— 
Eliason, 1920, p. 24 (part)—Chamberlin, 1920, p. 11——Hartman, 1944a, 
pp. 335, 338; 1948, p. 19.—Zatsepin, 1948, p. 111, pl. 29, fig. 2. 


Description.—Length 16-285 mm., width without setae 1-9 mm. 
(length up to 450 mm.—Ditlevsen, 1917). Segments very numerous. 
Body elongate, large, robust, linear, flattened, attenuate posteriorly. 
Prostomium cordiform, notched posteriorly with an occipital tubercle 
in the notch, with four short subulate frontal antennae, with small, 
paired, lateral, retractile, nuchal knobs just anterior to the first pair 
of tentacular cirri. Tentacular cirri unequal; two pairs short, two 
pairs longer, extending to segments 9-11. Dorsal cirri of median 
region up to two times longer than wide, subquadrangular, with distal 
ends truncate; ventral cirri oval, with asymmetrical acuminate tip. 
Anterior part of extended proboscis transversely rugose, crowned 
with 17 papillae; basal part with 12 longitudinal rows of small, oval 
papillae (6 rows on each side), with 10-20 papillae per row. Conor: 
In life and in alcohol: Body with bluish iridescence, usually heavily 
pigmented dark greenish brown or deep reddish blue, with dorsal and 
ventral cirri tan to dark brown; on some, body not so darkly pig- 
mented, with cirri only partly pigmented. 


MARINE POLYCHAETE WORMS—PETTIBONE 237 


Ficure 27.—Phyllodocidae: a, Eteone, dorsal view anterior end; b, Mystides borealis, 
dorsal view anterior end; c, Ewmida minuta, dorsal view anterior end; d, Phyllodoce 
groenlandica, dorsal view anterior end; ¢, Eteone barbata, parapodium; f, Eteone spets- 
bergensis, parapodium; g, Eteone flava, parapodium; h, Eteone longa, parapodium; 1, 
Phyllodoce groenlandica, parapodium. (For explanation of symbols, see p. 210.) 


238 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Remarks.——Some smaller specimens were at first placed under P. 
citrina Malmgren, since the papillae on the proboscis were fewer than 
normal, four rows on each side, with 3-5 papillae per row. An inter- 
mediate-sized specimen from Canoe Bay, Alaska (Hartman, 1948, as 
P. citrina) has five rows on each side, with as many as 6 papillae per 
row. It appears that the papillae are added on gradually and that 
only in the fully developed individual is the full complement of 6 
rows of papillae on each side developed; the number per row is varia- 
ble, ranging from 10 (with some spaces as if some had dropped off) 
to 20. 

New records —Arctic ALASKA: Eluitkak Pass, Elson Lagoon near 
Point Barrow; Point Barrow base, washed ashore; off Point Barrow 
base, up to 12.1 miles from shore, 1.7—123.5 fms., on bottoms of mud, 
sandy-mud, mass of worm tubes, and various combinations of mud, 
eravel, stones, pebbles, rocks, large perforated rocks, shells; in screen 
trap on bottom (23 stations, 74 specimens). Wrst GREENLAND: 
Vaigat, Disko Island, mud, Bartlett, 1937. East Coast Norra 
America: Off Labrador, 70 fms., Blue Dolphin Expedition, 1950; off 
Maine, Massachusetts, intertidal to 42 fms., U. S. Fish Commission. 
West Coast Nortu America: Washington Sound, Pettibone. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Franz 
Josef Land, Barents Sea, Novaya Zemlya, Kara Sea; Also Iceland, 
Faroes, Norway to English Channel; Hudson Bay to Massachusetts; 
Bering Sea to Washington; north Japan Sea. In low water to 800 
fathoms. 


Genus Eumida Malmgren, 1865 
Eumida minuta (Ditlevsen, 1917) 
FIGURE 27, c 


Eulalia minutia Ditlevsen, 1917, p. 56, pl. 4, figs. 10, 12, 14.—? Gorbunov, 1946, 
. 38. 
Bulatia arctica Annenkova, 1946, pp. 185, 187, fig. 1, a-c. 
Deseription.—Length 1-8 mm.; width without setae 0.3-0.8 mm. 
Segments few (12-36). Body very small, slightly tapered anteriorly 
and posteriorly, flattened dorsoventrally. Prostomium semiglobular 
to bluntly conical, with two large eyes near posterior border, with 
antennae rather long, slender, filamentous, almost as long as pro- 
stomium. First tentacular achaetous segment not distinct dorsally, 
with first pair tentacular cirri lateral to prostomium at same level as 
the eyes (thus placed under Humida and not Eulalia Savigny); two 
pairs tentacular cirri on second segment (first setigerous); fourth 
tentacular cirri on third segment; tentecular cirri enlarged basally, 


MARINE POLYCHAETE WORMS—PETTIBONE 239 


with slender filamentous tips, the latter about as long as the enlarged 
basal part except for the ventral tentacular cirri on the second segment 
which are much shorter. Dorsal cirri oval or bluntly conical. 
Ventral cirri elongate oval. Anal cirri short, oval, 1.5 to 2 times 
longer than wide. Cotor: In alcohol: Greenish tan to brown. In 
life: Flesh ventrally, green and flesh dorsally, with salmon-colored 
eggs showing through. 

New records—Arctic AuasKa: Off Point Barrow base, 16.7—75.5 
fms., up to 8 miles from shore, on various combinations of gravel, 
small stones, rocks, bryozoans, hydroids, Sazicava shells (6 stations, 
46 specimens). 

Distribution.—Few scattered records in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait. In 16.7—75.5 fathoms. 


Family HEsionIDAE 


Prostomium usually with four eyes, two or three antennae, two 
biarticulate palps (fig. 28, a). Two to eight pairs tentacular cirri on 
one to four more or less distinct achaetous tentacular segments. 
Parapodia biramous or subbiramous; notopodia may be greatly re- 
duced, with notosetae simple or lacking; neurosetae compound (fig. 
28, 6). Dorsal cirri long, more or less distinctly articulated; ventral 
cirri shorter; two long anal cirri. Proboscis cylindrical, eversible, 
with marginal papillae, with or without horny jaws. 

Represented by a single species from Point Barrow. (The speci- 
mens from Point Barrow, referred to Castalia multipapillata Théel by 
Murdoch (1885), do not agree with this species nor with any other 
species of hesionid; the parapodia are biramous, both lobes being 
pointed, the notopodia are well developed, with capillary setae; 
neurosetae compound, with delicate tips; the proboscis has numerous 
small papillae; tentacular cirri four pairs; prostomium with two 
pairs eyes, pair biarticulate palps, antennae ? (missing or absent); 
the specimens are very small and in poor condition.) 


Genus Castalia Savigny, 1820; emend. Fauvel, 1923 


Castalia aphroditoides (Fabricius, 1780) 


Figure 28, a, b 


Nereis aphroditoides Fabricius, 1780, p. 296. 

Castalia arctica Malmgren, 1867, p. 32.—Annenkova, 1931, p. 203. 

Castalia fabricit Malmgren, 1867, p. 32.—Théel, 1879, p. 37, pl. 3, figs. 36, 37.— 
Berkeley and Berkeley, 1943, p. 130. 

Castalia aphroditoides Wirén, 1883, p. 401.—Augener, 1913, p. 260; 1928, p. 715.— 
Zatsepin, 1948, p. 114.—Wesenberg-Lund, 1950a, p. 13, pl. 3, fig. 14; 1950b, 
p. 44. 

Psammate aphroditoides Chamberlin, 1920, p. 13. 


240 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Description.—Length 2.5-18 mm., width without setae 0.7-2.5 mm. 
Segments 25-43. Body cylindrical. Prostomium rectangular, wider 
than long, widest anteriorly, notched posteriorly, with two pairs 
eyes in trapezoidal arrangement, anterior pair larger; palps on an- 
terolateral borders of prostomium; two filiform antennae medial to 
palps (without unpaired antenna); antennae subequal, about same 
length as prostomium. Six pairs tentacular cirri on three more or less 
distinct achaetous segments; tentacular cirri long, filiform, articulated, 
with prominent cirrophores. Without distinct notopodia (acicula in 
cirrophores of dorsal cirri only); without notosetae. Neuropodia 
well developed, with three diverging conical distal lobes: Lip anterior 
to setae with median and dorsal conical lobes; lip posterior to setae 
with rounded median and conical ventral lobes; neurosetae compound. 
Dorsal and anal cirri long, articulated. Proboscis barrel-shaped with 
10 soft papillae around opening (four dorsal to lateral grooves, three 
each on two ventrolateral folds), with a median ventral notch, with 
two thickened ventrolateral ridges (the so-called jaws) lateral to the 
ventral notch. Cotor: In alcohol: Without color, or slightly greenish 
dorsally (especially in anterior part), or brownish. In life: Orange- 
flesh, with greenish eggs inside, with red eyes. 

Remarks——Two young specimens found in plankton tow beneath 
ice February 28, 1950. 

New records—Arctic AtAsKA: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms., stony-mud; off Point Barrow base, 21—-75.5 
fms., up to 8 miles from shore, on bottoms of mud, stones, and various 
combinations of mud, stones, gravel, rocks, and from breaking apart 
foliaceous bryozoans (10 stations, 28 specimens). NorrHwexst 
GREENLAND: Walrus grounds, Murchison Sound, Bartlett, 1938. 
East Coast Nortu America: Off Labrador, 45 fms., Blue Dolphin 
Expedition, 1949. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Novaya Zemlya, Kara Sea. Also Bering Sea; Denmark; Hudson Bay, 
Labrador. In 2-75.5 fathoms. 


Family SYLLIDAE 


Body of small size. Prostomium with four eyes (sometimes six), 
two palps (may be reduced and fused), three antennae (figs. 28, 29). 
Tentacular segment achaetous, with one or two pairs tentacular cirri. 
Parapodia uniramous, with dorsal and ventral cirri (latter may be 
absent). Two anal cirri. Setae mostly compound, sometimes with 
additional simple setae. Proboscis eversible, armed or not with one 


—_ 


MARINE POLYCHAETE WORMS—PETTIBONE 241 


to several chitinous teeth. Reproduction direct (epigamy) or by 
stolons, sometimes having sexual dimorphism; with swimming capillary 
setae at maturity. 

Represented by 6 genera and 11 species. 


Key to the genera of Syllidae from Point Barrow 


1. Without ventral cirri. Palps reduced, united, turned under on ventral side 
of prostomium (except for forked palps of male stolon; fig. 29, a). Antennae 
and dorsal cirri smooth or faintly annulated. Reproduction by stolons; 
stem forms produce by transverse fission, singly or in chains, sexually 
dimorphic male and female forms with swimming setae___ ----_ Autolytus—2 

With ventral cirri (fig. 28, d). Palps better developed, free or fused___-___-_- 4 

2. Prostomium normal, with 4 dorsal eyes in trapezoidal arrangement, with 3 
antennae, short fused palps; 2 pairs tentacular cirri (fig. 29, a). Without 
swimming setae. Pharynx long, more or less sinuous, usually with a crown 
GMCC UMM Oe R ne So oe Stem form or stock of Autolytus (p. 242) 

Prostomium abnormal, with 4 large eyes (usually 2 dorsolateral and 2 ventro- 
lateral) ; 2-3 pairs tentacular cirri (fig. 29, c-f)._ Body divided into an anterior 
unmodified region, a middle region with long swimming setae and well- 
developed, paddlelike parapodia, and with or without a posterior unmodified 
fang. s Without pharynx. ............-.. Sexual stolons of Autolytus—3 

3. Prostomium with 3 antennae (very long median one and pair of small frontal 
ones) and pair of anterior bifurcate palps; 1 pair tentacular cirri very long, 
similar to median antennae (these may be the dorsal cirri of the first setiger 
or of asmall, knoblike achaetous lobe; fig. 29, e, f). Testes and sperm con- 
fined to anterior unmodified region. 

Male form of Autolytus (Polybostrichus) (p. 242) 

Prostomium with 3 subequal antennae, without palps; none of tentacular 
cirri especially elongate (fig. 29, c, d). Most of body filled with eggs; eggs 
carried in large sac on ventral surface (gestation). 

Female form of Autolytus (Sacconereis) (p. 242) 

4. Dorsal cirri distinctly moniliform throughout body (fig. 28, c). Palps separate 
for their entire length or fused basally. Reproduction direct or by single 
stolons. Sexual stolons (Chaetosyllis): Small prostomium, with 2 pairs 
eyes, pair short moniliform antennae; without tentacular cirri; 1 anterior 
unmodified segment, a middle region with long swimming setae, and few 


unmodified, posterior segmentss—_<=la= - 2244525224. Js. Syllis (p. 252) 
Dorsal cirri smooth or indistinctly articulate (may be more or less articulate 
anteriorly). Palps fused at base only or throughout_--_._.__--_--- .-- 5 


5. Palps fused for nearly their entire length (fig. 28, k-m). Body very small. 
Antennae and dorsal cirri short, not moniliform. One pair tentacular cirri 
(may be rudimentary). Reproduction generally direct, with swimming 
setae at maturity; eggs and larvae fixed to dorsal or ventral surface of 
SES TR saa hg pe ks pel 6 

Palps fused at base only (fig. 28, g). Body larger. Antennae and dorsal 
cirri longer, smooth (antennae and anterior dorsal cirri may be indistinctly 
articulate, especially distally). Two pairs tentacular cirri. Reproduction 
direct, with swimming setae at maturity__._........-.._.------------ 7 


242 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


6. Antennae and dorsal cirri swollen at base, tapering to narrow tip (fig. 28, m). 
Body and parapodia usually covered with adhesive papillae and fine, granular 


100724 1 5 tap My Nee Ng ea i dy apes | bs Sphaerosyllis (p. 255) 
Antennae and dorsal cirri clavate or conical (fig. 28, k, 1). Body and parapodia 
not: covered wath. papillaes: 25-03. 202 ee es Exogone (p. 257) 


7. Compound neurosetae with distal blades all rather short (fig. 28, 7). Pro- 
boscis with a distal and proximal row of soft papillae, with distal circular 
chitinous margin denticled (fig. 28, g, h)_-.-_-___--_____- Eusyllis (p. 259) 

Compound neurosetae with some distal blades elongate (fig. 28, 7). Proboscis 
with distal row of soft papillae only, with distal circular chitinous margin 
SHO Otis: Sigs 3. BA DME RAL 3 tO oy ACS Pionosyllis (p. 262) 


Genus Auiolytus Grube, 1850 


In working over the rather numerous specimens of Autolytus in 
the collections from Point Barrow, several things were revealed which ~ 
throw light on the confusion in this group caused by the formation of 
sexually dimorphic stolons. For correct description of any species, 
one should have the stem form, the sexual buds in the process of 
formation including the mature sexual buds attached to the stem 
form, and the mature male and female stolons separated from the 
stem. Very often, however, stem forms and the male and female 
stolons of one species have been described and given different names, 
while sexual stolons and stem forms of different species have been 
given thesame name. A. fallax Malmgren, described originally from 
specimens from Spitsbergen, is of particular interest. Malmgren 
described and figured the stem form with the head of a male stolon 
forming between setigers 13 and 14 and indicated that the sexual 
stolons were not known. A. prismaticus (Fabricius), described 
originally from specimens from Greenland, has a characteristic color 
pattern of three dark longitudinal bands (median and lateral at the 
level of the bases of the dorsal cirri). The sexual stolons have six 
unmodified setigers anterior,to’ the setigers with ‘swimming setae; the 
stolons show the characteristic color pattern of the stem form. In 
the absence of color, it would be difficult to distinguish A. prismaticus 
from certain other species of Autolytus. It apparently has been the 
usual practice to identify any sexual stolon of unknown connections 
that has six anterior unmodified setigers as A. prismaticus, and some- 
times to identify the male stolons as Polybostrichus longosetosus 
Oersted or A. longisetosus (Oersted). However, as found in the Point 
Barrow material, A. fallax is the most common species of Autolytus 
and it was found along with A. prismaticus and A. alexandri. The 
majority of the specimens of A. fallax were in the process of stolon 
formation, the head of the stolon being formed between setigers 13 


MARINE POLYCHAETE WORMS—PETTIBONE 243 


Figure 28.—Hesionidae: a, Castalia aphroditoides, dorsal view anterior end; b, same, 
parapodium, posterior view. Syllidae: c, Syllis fasctata, dorsal view anterior end; d, 
same, parapodium; ¢, same, neuroseta; f, Syllis cornuta, neurosetae, with short (1) and 
long (2) blades; g, Eusyllis blomstrandi, dorsal view anterior end, proboscis partially 
extended; h, same, frontal view extended proboscis showing distal row of papillae and 
denticled margin; 1, same, neuroseta; 7, Pionosyllis compacta, neurosetae, lower one (1) 
with shorter blade, upper one (2) with longer blade; k, Exogone dispar, dorsal view anterior 
end; /, Exogone naidina, dorsal view anterior end; m, Sphaerosyllis erinaceus, dorsal view 
anterior end. (For explanation of symbols, see p. 210.) 


944 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


and 14. Although no mature sexual buds were found attached to the 
stem form, the rudiments of the notopodia with the developing swim- 
ming setae could be seen, beginning on setiger 7. Male and female 
stolons with six anterior unmodified segments were found. Of these, 
two female stolons showed the characteristic banding of A. prismaticus 
(this banding was noted on all the developing sexual buds still attached 
to the stem forms of this species). The others did not show this 
banding and are considered to be the sexual stolons of A. fallax. A. 
fallax, then, along with A. prismaticus, has six anterior, unmodified 
setigers and many of the records of A. prismaticus in the literature 
will no doubt prove to be A. fallax. 

The stem form of A. fallax has been referred to A. prolifer (Miiller) 
by Augener (1913, 1928) and Wesenberg-Lund (1947), thus the stem 
form of A. fallax has been confused with A. prolifer and sexual forms 
with A. prismaticus. 

There appears to be a correlation in the place of formation of the 
sexual stolons in the stem form with the number of anterior, unmodi- 
fied setigers and the number of tentacular cirri in the sexual stolons. 
At least for a group of species, including A. prismaticus, A. fallax, A. 
cornutus A. Agassiz, and A. ornatus (Verrill, 1873), the stem forms 
produce sexual stolons singly, with the head of the stolon forming 
between setigers 13 and 14. Thestolon, then, consists of the segments 
of the stem posterior to setiger 13 (differing from A. prolifer where 
the sexual stolons are proliferated more posteriorly, often in chains of 
2 to 8). The posterior part develops gradually into the male or 
female sexual stolon and breaks off when mature. The anterior 13 
setigers of the stem regenerates another posterior end. The sexual 
stolons of these four species have six unmodified setigers anterior to 
the swimming setae and usually a more or less developed unmodified 
posterior region. The stolons have three pairs of tentacular cirri on 
the first two achaetous tentacular segments; the first two pairs on the 
first segment and the third pair on the second segment. The upper 
pair of tentacular cirri on the first segment are similar to the dorsal 
curi. The lower pair may be very short, about the length of the 
prostomium. The third pair of tentacular cirri in the female are 
similar to the dorsal cirri; in the male they are very long, similar to 
the median antenna. In addition, there are a pair of small achaetous 
lobes. In contrast, in A. prolifer the sexual stolons have three ante- 
rior unmodified setigers (sometimes two or four) and a poorly devel- 
oped posterior region; there is a single tentacular achaetous segment 
with two pairs of tentacular cirri (in the female both the upper and 
lower pairs are short; in the male the lower one is similar to the dorsal 
cirri and the upper one is very long, similar to the median antenna). 
In A. alexandri the sexual stolons have 14 (sometimes 13) anterior 


——T ee 


MARINE POLYCHAETE WORMS—PETTIBONE 245 


unmodified setigers and a well-developed posterior region; there is a 
single achaetous tentacular segment with two pairs of tentacular cirri; 
the first setigerous segment in the female is similar to the following 
segment, but in the male it has a very long pair of dorsal cirri similar 
to the median antenna. Thus the male stolons of the different species 
of Autolytus agree in having a very long pair of anterior cirri similar 
to the median antenna, but they may be developed as the upper pair 
of tentacular cirri of a single tentacular segment as in A. prolifer, as 
the dorsal cirri of the first setigerous segment as in A. alexandri, or as 
the third pair of tentacular cirri on the second tentacular segment 
just dorsal to a pair of achaetous lobes as in A. prismaticus. 

The number of setigers in the anterior, unmodified region of the 
sexual stolons appears to be a good taxonomic character for most 
species. That the number is somewhat variable in A. prolifer (usually 
three, sometimes two and four) does not mean that it is of no signifi- 
cance in other forms. Thus the specimens of Autolytus with six 
anterior unmodified setigers, which Dales (1951) referred to A. prolvfer, 
are no doubt another species. The description agrees well with that 
of A. cornutus. 

All three species of Autolytus from Point Barrow have the body 
linear elongate, attenuated posteriorly, flattened ventrally, arched 
dorsally. Prostomium with four eyes in trapezoidal arrangement, 
with palps fused, turned ventrally, exceeding the prostomium only 
slightly. Neuropodia with composite setae with blades short, biden- 
tate, and usually with a simple bayonette seta. 


Key to the species of Autolytus from Point Barrow 


1. Stem Form: Dorsal cirri irregular in length, at least some longer than body 
width, pigmented with reddish globules; body whitish, brownish, or trans- 
versely banded with reddish globules, 2 narrow bands per segment; with 
nuchal epaulettes on first 2 (to 4) setigers; sexual buds formed singly (?), 
the heads forming posterior to setiger 14. Smxuau Sroions: 14 setigers 
anterior to swimming setae; nuchal epaulettes on first 3 setigers; 2 pairs 
tentacular cirri, the upper one longer; in male, first pair dorsal cirri on first 
setiger very long, similar to median antenna; female stolons with 2-lobed 
EN CMe Renan = age Sn RR: te NM ie ere eh Pa See A. alexandri 

Stem Form: Dorsal cirri, except anterior 2 pairs, shorter than body width, 
not extending beyond or only slightly beyond the setae, subequal; sexual 
buds formed singly, the heads forming between setigers 13 and 14. SEXUAL 
Srotons: 6 setigers anterior to swimming setae; 3 pairs tentacular cirri; in 
male, first pair similar to dorsal cirri, with second short pair at its ventral 
base, third pair very long and stout (similar to median antenna) with small 
achaetous lobe at its ventral base (fig. 29, e, f); in the female, similar except 
both the first and third pairs are similar to the following dorsal cirri (fig. 
29, c, d); female stolons with 1-lobed egg sac_-_---------------------- 2 


246 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


2. Stem Form: Usually with 2 dorsolateral longitudinal black to brown bands 
just dorsal to bases of dorsal cirri (may be faint or absent); with short 
semicircular epaulettes on tentacular segment only. SexuaL STOLOoNs: 
Colorless or female stolon dusky, especially posteriorly; male stolon rusty 
brown, darkest on anterior region, especially laterally; epaulettes on 
tentacular segment only (fig. 29, c—f)..-..---.---2-.--.4---2s28 A. fallax 

Stem Form: With 3 longitudinal black bands, 1 medium and 2 lateral at same 
level as bases of dorsal cirri; medium band continuous throughout length of 
body, lateral ones may fade out in anterior fourth or be lacking; with semi- 
circular epaulettes on tentacular segment and part of first setiger (fig. 29, a). 
SexuaL Stotons: With 3 longitudinal pigmented bands, median one con- 
tinuous throughout most of body, lateral ones along bases of dorsal cirri 
in anterior unmodified region; with epaulettes on tentacular segment and 
DarivOl, bite SAb@er 2 crs. ko ok A. prismaticus 


Autolytus alexandri Malmgren, 1867 


Autolytus alecandri Malmgren, 1867, p. 37, pl. 7, fig. 39.—Verrill, 1881, p. 292, 
pl. 12, fig. 8—Chamberlin, 1920, p. 12.—Friedrich, 1939, p. 123.—Hartman, 
1942a, p. 13; 1944a, p. 338, pl. 18, fig. 11 (not pl. 13, fig. 2); 1945, p. 17, 
pl. 2, fig. 11. 

Stephanosyllis picta Verrill, 1874a, pp. 43, 132, pl. 4, fig. 6; 1874b, pp. 361, 362, 
pl. 4, fig. 1 (not Proceraea picta Ehlers, 1864). 

Stephanosyllis ornata Verrill, 1874a, p. 132 (not Proceraea ornata Verrill, 1873). 

Proceraea (Stephanosyllis) ornata Webster and Benedict, 1887, p. 724 (not 
Proceraea ornata Verrill, 1873). 

Autolytus verrilli Marenzeller, 1892, p. 416, pl. 19, fig. 4—Augener, 1928, p. 726.— 
Wesenberg-Lund, 1947, p. 33, figs. 14, 15; 1950a, p. 18; 1950b, p. 52; 1951, 
p. 52. 

Description —Stem form: Length 4-12 mm., width without setae 
0.6-0.8 mm., up to 35-64 segments. Antennae rather short, thick, 
subequal or median one up to twice the length of the lateral; tentac- 
ular cirri equal to or shorter than antennae, upper pair longer; 
first pair dorsal cirri as long as or longer than median antenna; rest 
of dorsal cirri rather irregular, unequal, at least some longer than body 
width, with cirrophores prominent. Distinct epaulettes on first two 
setigerous segments (or to end of setiger 4—Hartman, 1945). None 
showed sexual bud formation (sexual bud is formed between segments 
25 and 26—Hartman, 1945). Cotor: In alcohol: Colorless or trans- 
versely banded with reddish to brownish granules, may be two bands 
per segment. 

Female stolon (Sacconereis): A single specimen of 72 setigers, 
16 mm. long, 1.2 mm. wide without setae, 3.5 mm. wide with swimming 
setae, consisting of 14 anterior, 26 middle, and 32 posterior setigers. 
Prostomium rounded, with four large dorsal eyes, three antennae 
rather short, subequal; two pairs tentacular cirri of nearly same 
length as antennae; dorsal cirri of anterior part of unequal length but 
all much longer than body width. Thick epaulettes extending on 
first three setigers. The stolon, with swimming setae and filled with 


MARINE POLYCHAETE WORMS—PETTIBONE 247 


eggs, was dredged in 27 fathoms February 18, 1950. (A 2-lobed 
egg mass is carried ventrally—Wesenberg-Lund, 1947.) 

Male stolon (Polybostrichus): No specimens taken from Point 
Barrow. A specimen taken in plankton at Portage Bay, Alaska, was 
12 mm. long, 1.5 mm. wide without setae, 4 mm. wide including 
swimming setae, with 61 setigers consisting of 14 anterior, 36 median, 
and 11 posterior segments. Prostomium similar to A. fallax; two 
pairs tentacular cirri, upper pair longer. First pair dorsal cirri on 
first setiger very long, similar to median antenna (achaetous—Wesen- 
berg-Lund, 1947); dorsal cirri in anterior region long, unequal; in 
middle region short, about one-third length of elongated parapodial 
lobes. Large epaulettes on first three setigers. 

New records.—Arctic AtasKa: Off Point Barrow base, 16.7—123.5 
fms., up to 15 miles from shore, on bottoms of stones, worm tubes, and 
various combinations of mud, pebbles, gravel, stones, rocks, large 
perforated rocks, with bryozoans, hydroids, shells, and worm tubes 
(17 stations, 53 specimens). Wrst Coast Norra America: Albatross 
surface station, Portage Bay, Alaska; San Juan Channel, Washington 
Sound, 25 fms., Pettibone, 1940. East Coast Nortu America: Off 
Labrador, 30-40 fms., Blue Dolphin Expedition, 1949; Woods Hole 
region, Massachusetts, Pettibone, 1950. 

Distribution.—Scattered records in the Arctic: Arctic Alaska, 
Greenland, Spitsbergen, Franz Josef Land, Barents Sea. Also Iceland; 
Labrador to North Carolina; Bering Sea to Washington. In low 
water to 123.5 fathoms; sexual forms at surface. 


Autolytus fallax Malmgren, 1867 
FIGuRE 29, cf 


? Polybostrichus longosetosus Oersted, 1848, p. 183, pl. 5, figs. 62, 67, 71.—Tread- 
well, 1937, p. 28. 

Autolytus fallax Malmgren, 1867, p. 33, pl. 6, fig. 41.—Ditlevsen, 1929, p. 17. 

? Autolytus longisetosus Malmgren, 1867, p. 34, pl. 7, fig. 38. 

Autolytus prolifer Augener, 1913, p. 258; 1928, p. 724 .—Wesenberg-Lund, 1947, 
p. 19, figs. 8, 9 (part; not Nereis prolifera Miller, 1788). 

Autolytus peaoincls Wesenberg-Lund, 1947, p. 24 (part; not Nereis prismatica 
Fabricius, 1780). 

Description —Stem form: Length 3-18 mm., width without setae 
0.3-0.7 mm., up to 45-78 segments. Median antenna and first pair 
dorsal cirri very long; lateral antennae and upper tentacular cirri 
about half as long; lower pair tentacular cirri and second pair dorsal 
cirri shorter, about twice the length of the following cirri; rest of dorsal 
cirri short, less than half the body width and extending only slightly 
beyond the setae, subequal. Short epaulettes (a pair of shallow, semi- 
circular raised areas) on tentacular segment only. Of 68 specimens of 
the stem form, 45 showed bud formation of the sexual stolons, the 


IAS PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


head of the bud forming between setigers 13 and 14; 6 were broken 
after setiger 13, 5 consisted of 13 setigers plus a definite posterior 
growing zone, and 12 consisted of 36 setigers or less and showed no 
sign of bud formation. Of the 45 specimens showing bud formation, 
18 were young buds (sex?) showing four eyes and rudiments of head 
appendages; 17 were developing male stolons as indicated especially 
by the developing forked palps (at first, one part of fork about one- 
half the length of the other), large median antenna, and large dorsal 
tentacular cirri; the male buds consisted of a variable number of 
setigers (14-26 plus a definite growing zone, or 32-42 and tapered 
gradually to an attenuated posterior end); 10 were developing female 
buds as indicated by the developing three subequal antennae and the 
absence of forked palps; the female buds consisted of 40-65 setigers. 
Cotor: In alcohol: Two dorsolateral, longitudinal dark bands just 
dorsal to bases of dorsal cirri (almost all specimens showed at least 
faint bands). In life: Anterior part pale yellowish with dark dorso- 
lateral bands, posterior part orange. TusEs: Thin, tough, transparent, 
on hydroids as Lafoeina maxima Levinsen; either on the surface or 
surrounded by the sessile hydrothecae of the hydroid colony. 

Female stolon (Sacconereis): Length 5-15 mm., width without 
setae 0.5-1 mm., width including swimming setae 3.5-4 mm., up to 
26-51 segments. Body widest in median region, tapering slightly 
anteriorly and more so posteriorly, forming an attenuated tail region; 
divided into three regions composed of 6 anterior unmodified setigers, 
13-18 modified setigers with long, iridescent swimming setae, and 
3-27 posterior unmodified setigers. Prostomium short, broad, one 
pair eyes dorsolateral, one pair larger ones ventrolateral, both with 
lenses; antennae long, stout, subequal or median one slightly longer 
than lateral pair. Upper two pairs tentacular cirri similar to dorsal 
cirri of following segments, with a third pair of short, ventral, tentacu- 
lar cirri (about length of prostomium) and a pair of small, ventral, 
achaetous lobes. Dorsal cirri about as long as body width. Pair 
of shallow semicircular epaulettes on tentacular segment only. Pyri- 
form-shaped egg sac carried on ventral surface in region of setigers 
10-18, the egg mass flattened or slightly concave on side toward body, 
the narrower end of lobe anterior, the eggs orange-colored (in life). 
Cotor: In alcohol: Colorless, transparent, may be dusky posteriorly 
or somewhat dusky throughout. Two female stolons with eggs in- 
side body dredged in 75 fms. October 11, 1949; females with egg sacs 
taken from vertical plankton hauls in 13 fathoms March 29, April 
15, May 17, 1950; female with egg sac containing developing embryos 
dredged in 5 fathoms January 25, 1950. 

Male stolons (Polybostrichus): Length 7-9 mm., width without 


setae 0.8-1 mm., width including swimming setae 2 mm., 43-66 seg- 


MARINE POLYCHAETE WORMS—PETTIBONE 249 


ments, with body regions of 6 anterior, 27-30 middle, 9-10 posterior 
setigers, tapering gradually posteriorly. Prostomium short, wide, 
with a pair of eyes dorsolateral, a larger pair ventrolateral, all with 
lenses; palps bifurcated about basal third, the two branches subequal; 
very long, stout median antenna and very small lateral antennae 
anterior and medial to dorsal pair of eyes. Three pairs tentacular 
cirri with anterior dorsal pair similar to dorsal cirri, anterior ventral 
pair shorter than prostomial length, posterior dorsal pair very large, 
similar to median antenna, with a pair of achaetous knobs at their 
ventral bases. Dorsal cirri of anterior setigers about as long as body 
width, those of modified region about as long as the elongated, paddle- 
like parapodial lobes, those of posterior, taillike region short, digitiform. 
Shallow semicircular epaulettes on tentacular segment only. Cotor: 
In alcohol: Rusty brown, darkest on anterior part or colorless except 
anterior part. In life: Yellowish green. Taken in plankton, near shore, 
September 10, 1949; dredged in 27 fathoms February 18, 1950. 

Remarks.—As referred to under the systematic discussion of 
Autolytus, the stem form of A. fallax has been incorrectly referred to 
A. prolifer by Augener (1913, 1928) and Wesenberg-Lund (1947, 1950). 
The two may be distinguished by the different positions of the sexual 
buds, the greater length of the dorsal cirri in A. prolifer, and the color 
markings. The sexual stolons of A. fallax have been confused with 
A. prismaticus (see discussion above). 

New records —Arctic Auaska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 5-6.6 fms., mud, in tubes associated with hydroid 
Lafoéina maxima Levinsen; Point Barrow base, washed ashore, on 
same hydroid; off Point Barrow base, 5-75.5 fms., up to 8 miles 
from shore, on bottoms of mud, stones, and various combinations of 
mud, gravel, stones, rocks, large perforated rocks, with bryozoans, 
hydroids including Lafoéina maxima, and shells (16 stations, 85 
specimens); sexual stages from plankton near shore to 75.5 fms. (7 
stations, 22 specimens). Cape Smyth, Point Barrow Expedition, 
winter, 1883 (1 male, 2 females with egg sacs). East Coast Norra 
America: Off Labrador, 25-60 fms., Blue Dolphin Expeditions, 1949, 
1951. 

Distribution —Widely distributed in the Arctic: Alaskan and 
Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, Novaya 
Zemlya. Also Faroes; Labrador. In low water to 75.5 fathoms; 
sexual stolons in plankton. 


Autolytus prismaticus (Fabricius, 1780) 
Figure 29, a, b 


Nereis prismatica ? Miller, 1776, p. 218.—Fabricius, 1780, p. 302; 1799, p. 177, 
pl. 4, figs. 17-20. 
? Polybostrichus longosetosus Oersted, 1843, p. 183, pl. 5, figs. 62, 67, 71. 


250 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


? Autolytus longisetosus Malmgren, 1867, p. 34, pl. 7, fig. 38.—Verrill, 1881, p. 
292, pl. 12, fig. 10.—Moore, 1902, p. 274; 1909b, p. 134. 

Autolytus incertus Malmgren, 1867, p. 35, pl. 6, fig. 40. 

Proceraea gracilis Verrill, 1874a, pp. 48, 132, pl. 5, fig. 1; 1874b, pp. 361, 362, 
370, 379, pl. 3, fig. 2—Webster and Benedict, 1887, p. 723. 

Autolytus prismaticus Marenzeller, 1892, p. 420.—Chamberlin, 1920, p. 12.— 
Augener, 1928, p. 725.—Annenkova, 1934, p. 322; 1938, p. 156.—Not 
Berkeley and Berkeley, 1938, p. 48; 1948, p. 68.—Gorbunov, 1946, p. 38.— 
Wesenberg-Lund, 1947, p. 24, figs. 10-12 (part) —? Hartman and Reish, 
1950, p. 13. 

Autolytus trilineatus Berkeley and Berkeley, 1945, p. 318, fig. 1, a-b; 1948, p. 
69, fig. 100. 


Description.—Stem form: Length 5-24 mm., width without setae 
0.7-1 mm., up to 52-92 segments. Proportion of anterior appendages 
and dorsal cirri similar to A. fallax. Distinct epaulettes on tentacular 
segment and usually extending also on at least part of first setigerous 
segment. Of 7 specimens, 2 were without signs of bud formation, 
although the body was constricted more than normally between 
setigers 13 and 14; 4 showed sexual bud formation with the head 
developing between setigers 13 and 14; and 1 had 13 setigers with a 
newly regenerating posterior end. Co or: In alcohol: Creamy white 
with three conspicuous, longitudinal, black or dark bands, the median 
one broader, less dense, continuous throughout body, the dorsolateral 
ones narrower, darker, at level of bases of dorsal cirri, often confined 
to anterior fourth or half of body. 

Female stolon (Sacconereis): A specimen of 58 setigers, 9 mm. long, 
1 mm. wide without setae, 3 mm. wide with swimming setae, with 
body regions of 6 anterior, 18 middle, and 34 posterior setigers. 
Epaulettes somewhat triangular, extending on tentacular and first 
setigerous segments. Otherwise as in A. fallax. Co tor: In alcohol: 
Shows the same characteristic pigmentation as the stem form, three 
longitudinal, black pigmented bands. Two female stolons with 
swimming setae and eggs massed inside body dredged in 5 and 6 
fathoms August 6, 30, 1948. Male stolons (Polybostrichus): no 
specimens taken. 

Remarks.—Polybostrichus longosetosus Oersted, the male stolon 
(this might well be the male stolon of A. fallax, see systematic dis- 
cussion above), Autolytus incertus Malmgren, the female stolon, and 
Proceraea gracilis Verrill, the stem form, have been referred previously 
by Marenzeller (1892) to A. prismaticus. A. trilineatus Berkeley and 
Berkeley is herein referred to A. prismaticus; the description, including 
the characteristic three longitudinal bands, is in agreement. The 
record of A. prismaticus by the Berkeleys (1938, 1948) is doubted 
since the stem form agrees with A. cornutus or A. prolifer but not A. 
prismaticus; the sexual forms agree with A. cornutus or A. prismaticus 
in the number of six anterior unmodified setigers; they agree with A. 


MARINE POLYCHAETE WORMS—PETTIBONE 251 


F 


Ficure 29.—Syllidae: a, Autolytus prismaticus, stem form, dorsal view anterior end; 3 
same, parapodium; c, Autolytus fallax, female (Sacconereis) form, dorsal view anterior 
end; d, same, ventral view anterior end; ¢, Autolytus fallax, male (Polybostrichus) form, 


dorsal view anterior end; f, same, ventral view anterior end. (For explanation of symbols, 
see p. 210.) 


261112—54—_4 


252 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


cornutus as to size, about one-third the size of the sexual stage of A. 
prismaticus, and in the absence of semicircular epaulettes (specimens 
examined through the kindness of the Berkeleys). The record of A. 
prismaticus from Oregon by Hartman and Reish (1950) is doubted 
since it is based on male stolons only. 

New records.—Axrctic AuasKka: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms., stony-mud; off Point Barrow base, 5-70 fms., 
up to 7 miles from shore, on bottoms of stones, and various combina- 
tions of mud, gravel, stones (7 stations, 11 specimens). East Coast 
Nortrs America: Lagoon Pond, Martha’s Vineyard, Massachusetts, 
pile scrapings, Pettibone, 1951. 

Distribution.— Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Barents Sea, Novaya 
Zemlya. Also Iceland; Labrador to Massachusetts; Bering Sea to 
British Columbia (? Oregon); north Japan Sea. In low water to 267 
fathoms; sexual forms at surface. 


Genus Syllis Savigny, 1818 


Both species represented have the body elongate, slender, with 
numerous segments, flattened ventrally, arched dorsally, tapering 
slightly anteriorly and gradually posteriorly. Prostomium oval, wider 
than long; two palps wider basally, well separated except where their 
inner basal sides are fused (fig. 28, c). Dorsal cirri distinctly monili- 
form throughout body, alternately slightly longer and shorter. Ven- 
tral cirri digitiform, slightly shorter or longer than the parapodial 
lobes (fig. 28, d). Anterior end of extended proboscis with a smooth, 
chitinous rim, a single large, grayish dorsal tooth, with a ring of short, 
thick papillae, with a ring of 10 shorter papillae more basally. Male 
and female stolons or epitokous forms (Chaetosyllis Malmgren, 1867) 
developed from the posterior segments of the body; when fully 
developed consisting of small bilobed (or tetralobed) bulbous “head”’ 
with two pairs of eyes, a dorsal pair and a larger ventral pair, with 
a pair of short moniliform antennae; the first setigerous segment with 
neurosetae only, followed by a variable number of segments with 
additional long capillary swimming setae and usually a few unmodified 
posterior segments; stolons greatly distended with developing sex 
products; neurosetae, dorsal, ventral and anal cirri similar to those 
of stem form. 


Key to the species of Syllis from Point Barrow 


1. Compound neurosetae with short and long distal blades (fig. 28, f). Acicula 
of parapodia not particularly enlarged or protruding. Body segments very 
shortest EL) seas dled pelo thes aaieatrte. wehbe, lee S. cornuta 

Compound neurosetae with terminal blades differing very slightly in length 
(fig. 28, e). From 1 to 4 very large acicula usually protrude somewhat from 
parapodial tip (fig. 28, d). Body segments longer______-_____- S. fasciata 


MARINE POLYCHAETE WORMS—PETTIBONE 253 


Syllis (Ehlersia) cornuta Rathke, 1843 
FIGURE 28, f 


Syllis (Ehlersia) cornuta Rathke, 1843, p. 164, pl. 7, fig. 12—Malmgren, 1867, 
p. 43, pl. 7, fig. 45.—Southern, 1914, p. 37.—Fauvel, 1923, p. 267, fig. 100, 
g-i.—Augener, 1928, p. 718.—Monro, 1933, p. 34; 1937, p. 273; 1939b, p. 
387.—Wesenberg-Lund, 1947, p. 6; 1950a, p. 15; 1950b, p. 46; 1951, p. 36.— 
Zatsepin, 1948, p. 115, pl. 31, fig. 4. 

Chaetosyllis oerstedi Malmgren, 1867, p. 45, pl. 8, fig. 51 (epitokous form). 

Syllis oerstedi Théel, 1879, p. 40, pl. 2, fig. 33; pl. 3, figs. 34, 35.—Annenkova, 
1938, p. 150.—Gorbunov, 1946, p. 38. 

Syllis quaternaria Moore, 1906a, p. 352, fig. (epitokous form). 

Syllis alternata Moore, 1908, p. 321, fig. (p. 324); 1909a, p. 321.—Annenkova, 
1938, p. 148.—Rioja, 1941, p. 691, pl. 3, figs. 1-9.—Berkeley and Berkeley, 
1948, p. 77, fig. 115. 

Syllis (Ehlersia) heterochaeta Moore, 1909a, p. 322, pl. 15, figs. 1-4.—Annenkova, 
1938, p. 148.—Rioja, 1941, p. 694, pl. 4, figs. 7-10; Berkeley and Berkeley, 
1948, p. 76, fig. 113. 

Ehlersia cornuta Hartman, 1945, p. 15. 

Typosyllis alternata Hartman, 1948, p. 21. 

Description.—Stem form: Length 9-45 mm., width without setae 
1-1.2 mm. Segments very short. Prostomium with two pairs of eyes, 
anterior pair larger, crescentric, with or without pair of small ocular 
spots anterior to lateral antennae. Moniliform dorsal cirri with about 
14-22 articles (11-40). Neurosetae all compound, with longer and 
shorter blades, the longer ones 2—4 times longer than the shorter ones, 
with fine marginal fringe extending to near the tips (may give appear- 
ance of having a bidentate tip with a subterminal tooth) ; tips of short 
blades slightly hooked; tips of long, thin blades slightly hooked or 
with a slight knob (worn down?). In posterior segments, terminal 
blades of neurosetae not as long as in anterior segments. One speci- 
men with developing stolon of 41 segments beginning on segment 
75 of stem form. Cotor: In alcohol: (1) anterior segments dusky, 
then two narrow, reddish brown, transverse bands per segment on 
anterior and middle part of each segment; (2) rusty brown, with two 
darker bands per segment on part of body; (3) colorless. 

Male and female stolons (Chaetosyllis): Length 8-19 mm., width 
without setae 1-1.2 mm., width including setae 4-5 mm., segments 
30-70 (1 unmodified anterior segment, 24-62 modified segments with 
swimming setae, 0-7 unmodified posterior segments); dorsal cirri 
alternately longer and shorter, with 11-18 articles (14-25—Moore, 
1906); capillary setae much longer than the dorsal cirri; neurosetae as 
in posterior segments of stem form (distal blades not as long as in 
anterior region of stem form) ; some segments showed the characteristic 
banding of two narrow bands per segment. 


954 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Remarks.—The types of Syllis alternata Moore from Alaska and 
S. heterochaeta Moore from California were examined and are herein 
referred to S. cornuta (suggested previously by Monro (1933) for S. 
heterochaeta). The description of S. quaternaria Moore (1906), the 
epitokous form taken at the surface, Point Barrow, Alaska, agrees 
with that of the stolons of S. cornuta collected by Dr. MacGinitie in 
a plankton haul. 

New records—Anrctic ALtaska: Stem form: Off Point Barrow base, 
18.3-123.5 fms., up to 15 miles from shore, on bottoms of stones, mass 
of worm tubes, and various combinations of mud, pebbles, stones, 
gravel, rocks, large perforated rocks, and worm tubes (17 stations, 97 
specimens). Sexual stolons: Off Point Barrow base, 1.6 miles from 
shore, vertical plankton haul of 13 fathoms through hole in ice, March 
29, April 15, 1950 (12 specimens). Wersr Coast NortH AMERICA: 
Strait of Juan de Fuca and Puget Sound, Washington, 30-107 fms., 
Pettibone. East Coast Norton America: Off Martha’s Vineyard, 
86, 134, and 146 fms.; off Salem, Massachusetts, 35 fms., U. S. Fish 
Commission. 

Distribution.—Cosmopolitan. Widely distributed in the Arctic: 
Siberian and Alaskan Arctic, Greenland, Spitsbergen, Novaya Zemlya. 
Also Iceland, Norway to Madeira, Mediterranean; Red Sea, South 
Arabian coast, Persian Gulf; Indian Ocean (Zanzibar); off South 
Africa; Maine to North Carolina; north Japan Sea, Alaska to Panamé; 
South Pacific (Marquesas, Tahiti). In low water to 1,400 fathoms; 
sexual stolons in plankton. 


Syllis (Typosyllis) fasciata Malmgren, 1867 
FIGURE 28, c-e 


Syllis (Typosyllis) fasciata Malmgren, 1867, p. 43, pl. 7, fig. 47; pl. 8, fig. 52.— 
Augener, 1928, p. 719.—Fauvel, 1934b, p. 304.—Annenkova, 1934, p. 322; 
1938, p. 150.—Gorbunov, 1946, p. 38.—Wesenberg-Lund, 1947, p. 10, fig. 
2,a; 1950a, p. 16; 1950b, p. 47; 1951, p. 37.—Berkeley and Berkeley, 1948, 
p. 74, figs. 109, 110.—Zatsepin, 1948, p. 115, pl. 31, fig. 2. 

Syllis armillaris Treadwell, 1937, p. 28 (not Nereis armillaris Miiller, 1776; in 
USNM). 

Description.—Stem form: Length 8-75 mm., width without setae 
0.7-3 mm. Many small ones present; many broken ones and a 
number with regenerating posterior ends. Segments not as short as 
in S. cornuta. Prostomium with two pairs of eyes, anterior pair 
larger, crescentric. Moniliform dorsal cirri with about 24 articles 
(20-40; 12-17 in young specimens). Parapodia each with one to 
four large, pointed acicula usually protruding beyond distal tips of 
setal lobes. Neurosetae all compound, with terminal blades rather 
short, hooked, with tips entire; some setae may have blades broken 


MARINE POLYCHAETE WORMS—PETTIBONE 255 


off (not to be confused with simple setae). Cotor: In alcohol: (1) 
pigmented with reddish-brown, one wide band per segment (darker 
on posterior part of band); more posteriorly two bands per segment, 
lighter anterior and darker posterior ones; then one band per segment; 
then without color (when mature, banded color pattern on the seg- 
ments of the developing sexual stolon); (2) colorless, particularly in 
small specimens; (3) uniform brown mottling on anterior third of 
body (specimens from Washington). In life: Reddish rusty brown 
bands, one wide band or two narrower bands per segment, especially 
on the anterior fourth and posterior fourth (latter a developing stolon, 
with wide, orange-red bands). One specimen with developing stolon 
beginning on segment 71. 

Sexual stolons (Chaetosyllis): Two male stolons, 34-37 segments 
(one unmodified anterior segment, 31-33 modified segments with 
swimming setae, 2 or 3 small, unmodified posterior segments); long 
swimming notosetae subequal to dorsal cirri; dorsal cirri of first 
unmodified segment broken off at cirrophores; darkly pigmented 
reddish brown dorsally and ventrally. 

New Records.—Arctic Atasxa: Stem form: Off Point Barrow base, 
18.3-123.5 fms., up to 15 miles from shore, on bottoms of mud, stones, 
worm tubes, from breaking rocks and bryozoans, from interstices 
between pebbles and gravel covering tunicate, Molgula sp., from 
various combinations of mud, pebbles, stones, gravel, rocks, large 
perforated rocks, worm tubes, and shells (21 stations, 206 specimens). 
Male stolons: Off Point Barrow base, 1.6 miles from shore, vertical 
plankton haul of 13 fms. through hole in ice (March 29, 1950, 2 speci- 
mens). West Coast Norta America: Washington Sound, 17-21 
fms., Pettibone. Canapian Arctic: Center Foxe Basin, 25-31 fms., 
Bartlett, 1927. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Greenland, Jan Mayen, Spitsbergen, 
Franz Josef Land, Novaya Zemlya, Kara Sea. Also Iceland, Faroes; 
Bering Sea to southern California; north Japan Sea, China. In low 
water to 378 fathoms; sexual stolons in plankton. 


Genus Sphaerosyllis Claparéde, 1863 
Represented by a single species from Point Barrow. 
Sphaerosyllis erinaceus Claparéde, 1863 


FIGURE 28, m 


Sphaerosyllis erinaceus Claparéde, 1863, p. 45, pl. 13, fig. 38 (fide Fauvel).— 
Southern, 1914, p. 20.—Fauvel, 1923, p. 302, fig. 115, q-r.—Annenkova, 
1934, p. 322; 1938, p. 153.—? Rioja, 1943, p. 211, figs. 1-6.—Gorbunov, 1946, 
p. 38.—Zatsepin, 1948, p. 116, pl. 31, fig. 7. 


256 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Sphaerosyllis latipalpis Levinsen, 1882, p. 244——Augener, 1928, p. 722, pl. 11, 
fig. 3; 1939, p. 140.—Wesenberg-Lund, 1947, p. 138, fig. 4; 1951, p. 38. 

Sphaerosyllis longicauda Webster and Benedict, 1887, p. 720, pl. 3, figs. 35-39.— 
Eliason, 1920, p. 11, fig. 3. 

Oophylax minuta Treadwell, 1937, p. 29, figs. 6, 7. 

Brania sp. Hartman, 1944a, pl. 24, figs. 1, 2. 

Description.—Length 2-4.5 mm., width without setae 0.3-0.5 mm., 
segments 22-37. Body linear, tapering slightly anteriorly and 
posteriorly, oval in cross section. Body, including parapodia, covered 
with small papillae and incrusted with fine, granular material. Pro- 
stomium subrectangular, wider than long, with six black eyes, four 
larger ones (each with a lens, arranged in transverse line or slightly 
concave are near middle of prostomium) and two small anterior ones. 
Antennae subequal, bulbous basally, narrower distally. Paired 
antennae lateral to anterior eye spots; median antenna more posterior, 
nearly in line with the four larger eyes. Palps short, wide, rounded 
anteriorly; they may project more anteriorly, extending about the 
same length as the prostomium, with basal halves fused and distal 
halves separated by a narrow groove, or they may project more ven- 
trally, extending only about half the length of the prostomium, and 
fused except for a small anterior indentation; ventrally the palps 
are not fused although they may be closely approximated, extending 
back to the level of the four large eyes. Tentacular segment not 
distinctly set off from the prostomium, with a single pair of tentacular 
cirri which are similar to the antennae. Dorsal cirri short, slightly 
longer than parapodial lobes and shorter than the setae, inflated 
basally, tapering to narrow tips; they may be similar in shape through- 
out body or they may be only slightly inflated basally in middle and 
posterior regions. Dorsal cirri lacking on setiger 2. Ventral cirri 
digitiform, slightly shorter than parapodial lobes. Anal cirri larger 
than dorsal cirri; anal segment with several larger papillae. Para- 
podial tip usually with a larger papilla. Neurosetae consisting of a 
single long, simple, tapering, curved upper seta, the rest compound 
with distal blades long, subequal, entire, hooked, and finely pectinate. 
Colorless in alcohol. Female (21 fathoms, September 9, 1948) with 
large, rounded eggs (larger in diameter than parapodial length) fastened 
to dorsal surface rather irregularly between setigers 8-24, one to four 
per segment, attached between neuropodia and dorsal cirri and just 
medial to the dorsal cirri; another female (36 fathoms, October 6, 
1949) with large, oval-shaped developing larvae (developing setae 
visible) attached to the dorsal surface; swimming setae lacking (with 
swimming setae in male—Fauvel, 1923; swimming setae in both male 
and female beginning on setiger 8—Webster and Benedict, 1887; 
eggs on ventral base of feet—Wesenberg-Lund, 1947). 


——— 


MARINE POLYCHAETE WORMS—PETTIBONE 257 


Remarks—The type of Oophylax minuta Treadwell from Foxe 
Channel was examined and is herein referred to Sphaerosyllis erinaceus; 
although not mentioned in the original description, the specimen has a 
median antenna (seen with difficulty on the prepared slide); neuro- 
setae include a simple seta in addition to the compound ones, setiger 
2 lacks dorsal cirri. The type specimens of S. longicauda Webster and 
Benedict from Eastport, Maine, were examined also but they are 
unsatisfactory, being hardened, shrunken, and covered with crystals; 
the description is fairly complete and agrees with S. erinaceus as 
does the description by Augener of S. latipalpis Levinsen. The 
record by Rioja of S. erinaceus from México is questioned chiefly 
because of the position of the eyes. 

New records.—Arctic AtAska: Off Point Barrow base, 16.7-75.5 
fms., up to 8 miles from shore, on bottoms of stones and various 
combinations of gravel, stones, rocks, large perforated rocks, with 
bryozoans and hydroids (8 stations, 37 specimens). East Coast 
Norru America: Off Labrador, 25 fms., Blue Dolphin Expedition, 
1949. Vineyard Sound, Massachusetts, surface, U. S. Fish Commis- 
sion, August 3, 1881. 

Distribution.—Widely distributed in the Arctic: Siberian , Alaskan, 
and Canadian Arctic, West Greenland, Spitsbergen. Also Iceland, 
Denmark, Ireland, English Channel, Baltic; Labrador to Mas- 
sachusetts; Bering Sea; north Japan Sea; ? Mexico. In low water 
to 75.5 fathoms; sexual forms at surface. 


Genus Exogone Oersted, 1845 


Both species represented are very small, slender, linear-elongate, 
tapered slightly anteriorly and more so posteriorly, flattened ventrally, 
arched dorsally. Body colorless, translucent (in alcohol). Pro- 
stomium oval to subrectangular, arched somewhat anteriorly, much 
wider than long; four eyes, with lenses, in trapezoidal arrangement; 
palps fused dorsally into a rounded or triangular mass, with or without 
a slight anterior notch; palps separated ventrally by wide depression. 
Tentacular segment short, more or less distinct, the anterior part of 
which may form a fold covering the posterior part of the prostomium. 
Dorsal cirri shorter than parapodial lobes, ovoid to somewhat conical. 
Ventral cirri digitiform, shorter than parapodial lobes. Neurosetae of 
three kinds: (1) upper one simple, slightly curved; (2) composite, 
with longer, slender, pointed blades; (3) composite, with short, hooked 
blades. Everted pharynx crowned with circle of papillae and single 
conical tooth. With swimming setae at maturity. Large eggs, 
embryos or young (up to five setigers, resembling the adult) attached 
to ventral surface of female. 


258 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Key to the species of Exogone from Point Barrow 


1. Antennae subequal. Tentacular cirri rudimentary, reduced to small knobs 
(fig. 28, 1). Dorsal cirri lacking on setiger 2. Upper compound setae with 
rather’ short pointed bladesi2_ 2! LU0.UL2_ £522. 21002) 2st eee E. naidina 

Median antenna more than 4 times length of lateral antennae (fig. 28, k). 
Tentacular cirri only slightly smaller than the lateral antennae or dorsal 
cirri. Dorsal cirri present on all setigerous segments. Upper compound 
setae with Jong pointed blades_...-—.-~-..+--.-..|-- - =e E. dispar 


Exogone naidina Oersted, 1845 


Figure 28, l 


Exogone naidina Oersted, 1845, p. 20, pl. 2.—Augener, 1939, p. 140.—Zatsepin, 
1948, p. 116, pl. 31, fig. 8. 

Exogone gemmifera Pagenstecher, 1862, p. 267, pl. 25; pl. 26, figs. 1, 2, 6-8.— 
McIntosh, 1908, p. 151, pl. 59, figs. 5, 6—Southern, 1914, p. 17.—Eliason, 
1920, p. 11.—Fauvel, 1923, p. 305, fig. 117, a-d.—Annenkova, 1934, p. 322; 
1938, p. 154.—Rioja, 1943, p. 223, figs. 38-46.—Gorbunov, 1946, p. 38.— 
Thorson, 1946, p. 35, fig. 8—Berkeley and Berkeley, 1948, p. 79, fig. 118.— 
Hartman and Reish, 1950, p. 13. 

Description.—Length 3.5-5 mm., width without setae 0.3 mm., 
segments 32-36. Palps form rounded mass, 1.4—2 times length of 
prostomium. Antennae shorter than palps, subequal or median one 
slightly longer, clavate, inserted anterior to eyes in almost a straight 
line. Tentacular cirri very small, reduced to small knobs. Dorsal 
cirri lacking on setiger 2. Neurosetae of each parapodium consisting 
of one simple, curved upper seta, one upper composite seta with 
longer pointed blade, and four or five composite setae with short, 
hooked blades. Female (21 fathoms, September 15, 1948) without 
special swimming setae, with large eggs attached to ventral surface 
(between segments 11-25, one to three per segment) medial to ventral 
cirri and between parapodia. Another female (57 fathoms, October 
11, 1949) with eggs on ventral surface of segments 13-21 (mostly 
four per segment); eggs large, nearly touching. (Swimming setae on 
females with internal ova, absent in females with ova attached to 
ventral surface, two eggs per segment—Thorson, 1946.) 

New records —Axrctic ALAska: Off Point Barrow base, 16.7—123.5 
fms., up to 16 miles from shore, on bottoms of rocks, stones, worm 
tubes, and various combinations of mud, gravel, stones, rocks, large 
perforated rocks, with worm tubes, bryozoans, hydroids; on female 
Hyas coarctatus alutaceus (14 stations, 82 specimens). 

Distribution —Alaskan and Siberian Arctic; Denmark, Ireland, 
Great Britain to Madeira, Mediterranean; Bering Sea to Mexico; 
north Japan Sea. In low water to 123.5 fathoms; sexual stages at 
surface. 


MARINE POLYCHAETE WORMS—PETTIBONE 259 


Exogone dispar (Webster, 1879) 
FIGurReE 28, k 


Paedophylax dispar Webster, 1879a, p. 223, pl. 4, fig. 49; pl. 5, figs. 50-55; 1879b, 
p. 110; 1886, p. 138.—Sumner, 1913, p. 615. 

Paedophylax longiceps Verrill, 1879, p. 170; 1881, p. 320, pl. 12, fig. 2; 1882, p. 
370.—Andrews, 1891, p. 281. 

Paedophylax longicirris Webster and Benedict, 1887, p. 722, pl. 3, figs. 46-50. 

Exogone lourei Berkeley and Berkeley, 1938, p. 44, figs. 6-12; 1948, p. 79, fig. 117.— 
Rioja, 1941, p. 703, pl. 3, figs. 14-21; 1943, p. 224. 

Exogone dispar Hartman, 1942a, p. 11; 1944a, p. 338, pl. 24, fig. 9; pl. 25, fig. 5; 
1945, p. 16, pl. 2, figs. 7, 9, 10; 1951, p. 40. 

Description.—Anterior ends of 3 specimens only (up to 8 mm. long, 
45 setigers—Berkeley and Berkeley, 1938). Palps form a conical 
mass, about twice length of prostomium. Antennae inserted between 
and slightly in front of eyes; median antenna longer, usually reaching 
nearly to end of palps (may be not much longer than prostomium), 
lateral antennae short, ovoid. Tentacular cirri slightly smaller than 
lateral antennae and dorsal cirri. Dorsal cirri present on all setigerous 
segments, including setiger 2. Neurosetae on each parapodium con- 
sisting of one simple curved upper seta (may have a delicate thread 
coming off subterminally in some specimens), one to four composite 
setae with long terminal capillary blades, and three to nine composite 
setae with short, hooked blades. 

Remarks.—The specimens were compared with the cotypes of 
Paedophylaz dispar from New Jersey. ‘The types of P. longicirris 
Webster and Benedict from Eastport, Maine, were examined also 
and the peculiar hairlike process on some of the simple setae was 
noted. This also occurs on some of the simple setae of the specimens 
from Point Barrow. P. longiceps Verrill has previously been referred 
to P. dispar by Sumner (1913). The description of L. lowre: Berkeley 
and Berkeley from British Columbia agrees with that of E. dispar. 

New records—Arctic AtasKa: Off Point Barrow base, 36-70 fms., 
up to 7.5 miles from shore, on bottoms of stones, gravel, large per- 
forated rocks (2 stations, 3specimens). Wrst Coast NortH AMERICA: 
Strait of Juan de Fuca, Washington, shore, Pettibone, 1940. East 
Coast NortH America: Woods Hole region, Massachusetts, Petti- 
bone, 1950, 1951. Maine, Massachusetts, New Jersey, low water to 
14 fms., U. S. Fish Commission. 

Distribution —aArctic Alaska to Mexico; Maine to southwestern 
Florida. In low water to 70 fathoms; sexual stages at surface. 


Genus Eusyllis Malmgren, 1867 


Both species represented are flattened ventrally, arched dorsally, 
tapering both anteriorly and posteriorly, very fragile, breaking up 


260 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


easily and losing their antennae and dorsal cirri readily. Prostomium 
subrectangular to suboval, wider than long; four eyes fairly large, in 
trapezoidal arrangement, the anterior pair larger; palps broad, thick, 
oval to squarish, fused at their bases only and well separated distally, 
and may be curled ventrally or longitudinally like flopping ears; 
lateral antennae about 1.5 times the prostomial length; median 
antenna about twice the length of the lateral antennae (fig. 28, g). 
Upper tentacular cirri as long as median antenna, ventral pair much 
shorter. Ventral cirri thick, oval, about same size and length as 
parapodial lobes or may be slightly shorter (first pair not enlarged, 
similar to the following). Neuropodia short and plump, with neuro- 
setae all composite except on the last setigers where there may be a 
simple, entire, upper seta and a simple, bidentate, lower seta; blades of 
composite setae rather short, hooked, distinctly bidentate, finely 
spinous on the cutting edge; shaft distally spinous (fig. 28, 2). At 
maturity, with long capillary swimming setae. Proboscis with two 
rows of soft papillae, a proximal and a distal row of 10 papillae per 
row; inside the distal row, a chitinous lining provided with a large, 
triangular, dorsal tooth and a denticled, circular margin (fig. 28, g, h). 


Key to the species of Eusyllis from Point Barrow 


1. Cirrophores of dorsal cirri not prominent. Tentacular segment with a wide 
crescent-shaped nuchal lobe covering the posterior part of the prostomium 


(fie 28 Hg) apes © be tae 6 oi ee a eee ee E. blomstrandi 
Cirrophores of dorsal cirri prominent, as long as the neuropodia. Tentacular 
segment short, without nuchal lobe__--_______-__--___=-=-= E. magnifica 


Eusyllis blomstrandi Malmgren, 1867 


FIGuRE 28, g-7 


Eusyllis blomstrandi Malmgren, 1867, p. 40, pl. 6, fig. 43.—Fauvel, 1923, p. 293, 
fig. 112,h-m.—Augener, 1928, p. 721.—Annenkova, 1938, p. 153, fig. 6.— 
Wesenberg-Lund, 1947, p. 11, fig. 3; 1950a, p. 16; 1950b, p. 48; 1951, p. 37.— 
Berkeley and Berkeley, 1948, p. 84, fig. 126. 

Eusyllis monilicornis Malmgren, 1867, p. 41, pl. 6, fig. 44.—Verrill, 1881, p. 319.— 
Zatsepin, 1948, p. 116, pl. 31, fig. 6,b. 

? Eusyllis phosphorea Hartman, 1942a, p. 7; 1944a, pp. 334, 338, pl. 25, fig. 4. 

Syllis monilicornis Théel, 1879, p. 41, pl. 3, fig. 39. 

Eusyllis tubifex McIntosh, 1908, p. 173, figs —Southern, 1914, p. 32 (not Syllis 
tubifex Gosse, 1855). 

Eusyllis bloemstrandi Friedrich, 1939, p. 122. 

Typosyllis collaris Hartman, 1948, p. 23, fig. 6,a-c. 


Description.—Length 7-32 mm., width without setae 0.8-1.2 mm., 
segments 50-124. Antennae, tentacular cirri, anterior dorsal cirri, 
and long anal cirri irregularly annulated, especially distally, more or 
less smooth basally; the more posterior dorsal cirri only indistinctly 
annulated or smooth. First few dorsal cirri may be longer than body 
width, especially the first pair which may be the longest appendage; 


: 


MARINE POLYCHAETE WORMS—PETTIBONE 261 


rest of dorsal cirri shorter than body width, tapering gradually. 
Tentacular segment with wide, short, crescent-shaped collar or nuchal 
fold covering posterior part of prostomium, may be inconspicuous 
and protruding or rather low, somewhat contracted. Composite 
setae similar throughout, with distal blades short, differing only 
slightly in length. Chitinous lining of proboscis light or dark gray 
or light amber-colored; dorsal tooth dark bluish gray; circular margin 
finely denticled. Epitokous female (Elson Lagoon, August 6, 1948) 
with large eggs massed in most of body segments, consisting of 16 
anterior unmodified setigers, 25 setigers with long swimming setae, 
and 5 unmodified setigers (Swimming setae begin on setiger 13— 
Wesenberg-Lund, 1947). Cotor: In alcohol: Colorless or tannish, 
slightly greenish anteriorly. In life: Orange or yellowish. May form 
definite hardened mucous tubes. Luminescent. 

Remarks.—The type of Typosyllis collaris Hartman (1948) was 
examined and is herein referred to E. blomstrandi; the dorsal cirri 
are not distinctly annulated throughout the body as they would be 
ina Typosyllis. There is nothing in Verrill’s description of E. phos- 
phorea, dredged from St. George’s Bank, to distinguish it from E. 
blomstrandi; according to Hartman (1942, p. 7) the type is unsatis- 
factory. 

New records —Arctic Ataska: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; Point Barrow base, washed ashore; off Point 
Barrow base, 13.3-123.5 fms., up to 15 miles from shore, on bottoms 
of stones, masses of worm tubes, and various combinations of mud, 
pebbles, gravel, stones, rocks, large perforated rocks, with worm tubes, 
bryozoans, hydroids, shells, and on female crab Hyas coarctatus alu- 
taceus (24 stations, 175 specimens). Brritne Sa: Robert White, 
1879. West Coast Nortu America: Washington and Puget Sounds, 
20-46 fms., Pettibone. Hast Coast Nortu America: Off Labrador, 
15-30 fms., Blue Dolphin Expeditions, 1949, 1951. 

Distribution.—Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, West Greenland, Spitsbergen, Franz Josef Land, 
Barents Sea, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Ire- 
land to Madeira, Mediterranean; Labrador to Maine; Bering Sea to 
Washington; north Japan Sea. In low water to 444 fathoms; epitokes 
at surface. 

Eusyllis magnifica (Moore, 1906) 


Pionosyllis magnifica Moore, 1906b, p. 2238, pl. 10, figs. 9-11; 1908, p. 325.— 
Annenkova, 1938, p. 152. 

Deseription.—Atokous specimen 52 mm. long, 2.5 mm. in greatest 
width without setae, 152 segments (in pieces, complete?). Epitokous 
specimen (18 fathoms, September 16, 1948) broken up and not quite 
complete, with larger eyes, consisting of 161 segments, 99 mm. long, 


262 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


4 mm. wide without setae, 9 mm. wide including long swimming setae. 
Two atokous specimens from Washington 43-48 mm. long, 1.5-2 mm. 
wide without setae. Median antenna, tentacular cirri and some of 
anterior dorsal cirri somewhat annulated, especially distally; rest of 
dorsal cirri smooth. Some of anterior dorsal cirri longer than body 
width; rest shorter than body width, subulate, thick basally, tapering 
distally, on prominent cirrophores which may be as long as parapodial 
lobes. Tentacular segment very short, without nuchal collar. Blades 
of neurosetae all rather short. In anterior parapodia, blades of upper 
neurosetae somewhat longer than the lower ones; on rest of parapodia, 
blades of neurosetae only slightly different in length. Chitinous lining 
of proboscis and dorsal tooth light to darker amber-colored or trans- 
parent; circular anterior border with coarse, triangular denticles (when 
transparent, difficult to see denticled border). Swimming setae begin 
on setiger 24 (on setiger 19 in specimen from Portage Bay, Alaska). 
Cotor: In alcohol: Flesh. 

Remarks.—The specimens were compared with the type of Piono- 
sylis magnifica Moore (1906) from Port Townsend, Washington. 
It is herein referred to Husyllis since the composite neurosetae have 
rather short blades which differ only slightly in length, the proboscis 
has a proximal and a distal row of papillae, and the anterior circular 
border is denticled (difficult to see when the chitinous lining is 
transparent). 

New records—Arctic ALaska: Off Point Barrow base, 18.3 fms., 
pebbles-mud (1 station, 2 specimens). Wurst Coast NortH AMERICA: 
Albatross surface station, Portage Bay, Alaska; Washington Sound, 
Pettibone. 

Distribution.—Arctic Alaska to Washington; north Japan Sea. In 
16 to 302.8 fathoms; epitokes at surface. 


Genus Pionosyilis Malmgren, 1867 
Pionosyllis compacta Malmgren, 1867 
FIGureE 28, 7 


Pionosyllis compacta Malmgren, 1867, p. 40, fig. 48.—Augener, 1928, p. 722.— 
Annenkova, 1934, p. 322. 

Description—Anterior ends only of two small specimens, up to 
1.5 mm. wide without setae (8-10 mm. long, 1 mm. wide—Malmgren, 
1867). Body flattened ventrally, arched dorsally, fragile, with seg- 
ments crowded. Prostomium subrectangular, wider than long; four 
eyes rather large, in trapezoidal arrangement; palps squarish, fused 
basally, widely separated distally. Tentacular segment without 
nuchal collar. Antennae, tentacular, and dorsal cirri smooth, without 
articulations, filiform to rather thick, breaking off readily; most of 


MARINE POLYCHAETE WORMS—PETTIBONE 263 


dorsal cirri shorter than the body width. Ventral cirri thick, pointed, 
almost as long as parapodial lobes. Neurosetae all composite, distal 
blades distinctly bidentate, finely spinous along cutting edge; blades 
of upper setae longer, becoming gradually shorter in lower part of 
bundle with upper ones about twice the length of lower ones. Proboscis 
with a distal row of 10 papillae; transparent chitinous lining with a 
translucent bluish-gray dorsal tooth, with distal rim smooth. Cotor: 
In alcohol: Irregularly dotted with black pigment both dorsally and 
ventrally. (Epitokous forms have capillary setae beginning on seg- 
ments 12-15—Malmegren, 1867.) 

New records—Arctic AuasKa: Off Point Barrow, 36-78 fms., up 
to 15 miles from shore, on bottoms of rocks, stones, and worm tubes 
(2 stations, 2 specimens). 

Distribution —Arctic Alaska; Spitsbergen; Bering Sea. In low 
water to 78 fathoms. 


Family NEREIDAE 


Prostomium distinct, with four eyes, two frontal antennae, two bi- 
articulated palps (fig. 30, @). Four pairs tentacular cirri; buccal or 
tentacular segment usually apodous and achaetous. Parapodia 
usually biramous (except first two pairs), with dorsal and ventral 
cylindrical cirri, two or three notopodial ligules and one neuropodial 
ligule (fig. 30, b, c). Setae usually compound spinigers and falcigers 
(fig. 30, d-g). Pair of anal cirri. Proboscis eversible, with pair of 
terminal, horny, faleate jaws and usually with horny denticles or 
paragnaths arranged in eight groups on two rings, a basal or oral 
ring and a distal or maxillary ring (fig. 30, A, 7). At maturity usually 
with a pelagic epitokous or heteronereis stage. 

Represented by a single genus and two species. 


Genus Nereis Cuvier, 1817 
Subgenus Nereis Linné, 1758 


Both species have the body smooth, elongate, cylindrical, at- 
tenuated posteriorly, with numerous segments. Prostomium of the 
typical nereid form. Tentacular segment achaetous, over twice the 
length of the following segments; tentacular cirri short, fairly uniform 
in length, the upper one only slightly longer than the others (may 
reach setiger 2). Parapodia biramous, with two subequal noto- 
podial ligules (remaining so throughout the body, the upper one not 
getting larger or longer than the lower one), a single neuropodial 
ligule ventral to the conical setigerous neuropodial lobe (fig. 30, }, ¢). 
Notosetae homogomph spinigers (fig. 30, g); in more posterior seg- 
ments with some homogomph falcigers with blunt asymmetrical 
reduced appendages (fig. 30, e). Neurosetae homogomph and 


264 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


heterogomph spinigers (fig. 30, f) and heterogomph falcigers with 
hooked end-pieces (fig. 30, d). Proboscis with paragnaths all 
chitinous, conical, separated in groups, occurring on both rings 
(fig. 30, h, 7). Distal ring of paragnaths: Area II (dorsolateral)— 
oblique group of two or three rows; area III (medioventral)—small 
transverse group; area IV (ventrolateral)—arched mass. Basal ring 
of paragnaths: Area V (mediodorsal)—0. At maturity, with epito- 
kous stage, eyes enlarged; 14-19 anterior prenatatory setigers; 
modified natatory setigers with parapodial lobes compressed, flat- 
tened anteroposteriorly, developing foliaceous lamellar plates, with 
numerous paddlelike swimming setae; in males, first seven dorsal 
cirri enlarged and club-shaped, first five or six ventral cirri modified, 
and dorsal cirri of natatory setigers crenulate on lower margin; in 
females, dorsal and ventral cirri not modified or crenulate. 


Key to the species of Nereis (Nereis) from Point Barrow 


1. Parapodial ligules short, thick, evenly rounded (fig. 30, 6). Body uniformly 
purplish or reddish brown, not banded. Paragnaths of distal ring: Area 
I (mediodorsal)—2 in tandem (rarely 1 or 3); paragnaths of basal ring: 
Area VI (dorsolateral)—4 in square or cross (rarely 3 or 5); areas VII- 
VIII (ventral)—1-2 irregular rows of larger paragnaths, with several rows 
of small ones, diminishing in size posteriorly (2-3 irregular rows of subequal 
paragnaths in subspecies occidentalis). Male epitokes with 16 and females 
with i”—19,prenatatory ; setigers: 2:2 22cu2 222)e8-.-15 4 N. (N.) pelagica 

Parapodial ligules triangular to conical, gradually tapering to a broad tip 
(fig. 30, c). Body transversely banded reddish brown or violet. Parag- 
naths of distal ring: Area I—O or 1; paragnaths of basal ring: Area VI— 
6-10 or more in oval mass; VIJ—-VIII—continuous row of larger parag- 
naths followed by a wide band of small subequal ones (fig. 30, A, 7). Male 
epitokes with 14 and females with 16 prenatatory setigers___.N. (N.) zonata 


Nereis (Nereis) pelagica Linné 1758 
Figure 30, a, 5 


Nereis pelagica Linné, 1758, p. 654.—Malmgren, 1867, p. 47, pl. 5, fig. 35.— 
Webster and Benedict, 1884, p. 718; 1887, p. 724.—Moore, 1903, p. 431.— 
McIntosh, 1908, p. 268, figs—Sumner, 1913, p. 619.—Southern, 1914, p. 
80.—Eliason, 1920, p. 29.—Chamberlin, 1920, p. 14.—Fauvel, 1923, p. 336, 
fig. 130, a-f—Augener, 1928, p. 712.—Monro, 1930, p. 106.—Annenkova, 
1934, p. 322; 1937, p. 162; 1938, p. 158.—Treadwell, 1937, p. 29.—Okuda, 
1939, p. 230.—Friedrich, 1939, p. 123.—Hartman, 1940, p. 225, pl. 35, fig. 
52; 1944a, p. 335; 1948, p. 26.—Berkeley and Berkeley, 1948, p. 130; 1948, 
p. 66, fig. 96, a-b.—Thorson, 1946, p. 64, fig. 29.—Zatsepin, 1948, p. 119, 
pl. 30, fig. 2—Wesenberg-Lund, 1950a, p. 19, pl. 6, figs. 26a, 27a; 1950b, 
p. 53; 195], p. 39. 

Heteronereis grandifolia Rathke, 1843, p. 155, pl. 7, figs. 13, 14.—Malmgren, 
1865, p. 108, pl. 11, fig. 15 (part—female heteronereid, not male). 

Nereis neonigripes Hartman, 1936b, p. 471, fig. 48. 

Nereis arctica Treadwell, 1937, p. 30. 

Nereis pelagica var. occidentalis Hartman, 1945, p. 20, pl. 4, figs. 1-6; 1951, p. 46. 

Nereis pelagica neonigripes Hartman and Reish, 1950, p. 17. 


MARINE POLYCHAETE WORMS—PETTIBONE 265 


Description —Length 60-155 mm., width 5-14 mm. See key. 
Cotor: In alcohol: Iridescent, uniformly purplish or reddish brown 
(narrow white intersegmental lines); ligules of parapodial lobes may 
be darkly pigmented (var. neonigripes); tips of dorsal cirri may be 
black. 

New records —Arctic ALASKA: Off Point Barrow base, up to 12.1 
miles from shore, 18-87 fms., on various combinations of stones, 
gravel, rocks, large perforated rocks (8 stations, 10 specimens). 
SPITSBERGEN: Spitsbergen Sea, U.S.S. Alliance, August, 1881, surface 
(heteronereid). Brrtna Sra: Albatross Sta. at Nikolski, Bering Island, 
1892, and Sta. 3496, 56°32’ N., 169°45’ W., 41 fms., 1893; St. George 
Island, Pribilofs, 1913, 1914, low tide and 30 fms., and St. Paul Island, 
Pribilofs, lagoon reef, 1915, Hanna; St. Paul Island, Wm. Palmer, 
1890. SourHwssterRN Ataska: Albatross Sta., Unalaska, Aleutians, 
1888; Akutan Pass, Chica Islands, 1872; Kiska Harbor, sandy- 
mud, 9-12 fms., 1873; Anchorage, Big Koniuji Islands, Shu- 
magin, 6-20 fms., sand and rocks; Port Moller, Alaska Peninsula, 
beach to 17 fms., sand, 1874; Killisnoo, 1897; Chiniak Bay, Kodiak 
Island, 1880; all collected by Dall. Gutr or Atasxa: Albatross Sta., 
at Observation Island, Cordova, Alaska, 1914. SouTHEASTERN 
Auaska: Sitka, 1932. Wzxst Coast NortH America: Albatross 
Sta. 89a, Denman, British Columbia, 1914, and Sta. 3466, off Wash- 
ington, 48°18’30’’ N., 123°22’ W., 56 fms., 1891. Washington and 
Puget Sounds, low tide to 166 fms. (very common), Pettibone. East 
Coast Norta America: Off Labrador, 7-60 fms., Blue Dolphin 
Expeditions, 1949, 1951; Woods Hole region, intertidal and dredged, 
Pettibone, 1950, 1951; Gulf of St. Lawrence, New Brunswick, Nova 
Scotia, Bay of Fundy, Maine, Massachusetts, Rhode Island, Con- 
necticut, Long Island Sound, shore to 250 fathoms and surface, U. S. 
Fish Commission. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Barents 
Sea, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Norway to 
Mediterranean; Hudson Bay to Long Island Sound; Bering Sea to 
Panamé; north Japan Sea, Japan; South Atlantic (Tristan da Cunha, 
Kerguelen, Magellan Straits). In low water to 609 fathoms; epitokes 
at surface. Variety occidentalis Hartman: North Carolina, Florida, 
Louisiana, Texas, Puerto Rico; intertidal. 


Nereis (Nereis) zonata Malmgren, 1867 
Figure 30, c, h, 7 


Heteronereis grandifolia Malmgren, 1865, p. 108, pl. 11, fig. 16 (part—male 
heteronereid, not female; not H. grandifolia Rathke, 1843). 

Nereis zonata Malmgren, 1867, p. 46, pl. 5, fig. 34.—Théel, 1879, p. 42.— Wirén, 
1883, p. 402.—Ditlevsen, 1911, p. 419, pl. 28, fig. 6; pl. 30, figs. 18, 22— 


266 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Fauvel, 1914, p. 177, pl. 14, figs. 1-17; 1923, p. 338, fig. 130, g—n (part).—_ 
Augener, 1928, p. 713.—Annenkova, 1931, p. 203; 1932, p. 134; 1934, p. 
322; 1937, p. 162; 1938, p. 158.—Friedrich, 1939, p. 123.—Berkeley and © 


Berkeley, 1943, p. 130.—Gorbunov, 1946, p. 38.—Zatsepin, 1948, p. 119, 


pl. 30, fig. 3—Hartman, 1948, p. 25.—Hartman and Reish, 1950, p. 17.— _ 


Wesenberg-Lund, 1950a, p. 20, pl. 6, figs. 26b, 27b; 1950b, p. 55; 1951, p. 41. 


Description.—Length 30-125 mm., width 7 mm. Seekey. Couor: 


In alcohol: Reddish or rusty brown or violet wide transverse bands; 
anterior fourth of segment without color. 


: 


Remarks.—Nereis procera Ehlers, 1868, originally described from | 


the Gulf of Georgia, British Columbia, has been referred to N. zonata— 


j 


by Fauvel, 1914. In N. procera the body is long, slender, extremely — 


attenuated posteriorly; the notopodial ligules of the anterior setigers 


are triangular, subequal; those of the median and posterior regions — 


have the upper notopodial ligules larger than the lower one (may be — 


‘ 


nearly twice the size); the paragnaths of areas VII and VIII are in a © 
wide, continuous area with no especially enlarged anterior row; male — 
epitokes with 51 and females with 59 prenatatory setigers. It may be | 
that the variety procera of Fauvel refers to a different species from that 


of N. procera Ehlers. 


New records—Arctic AuasKA: Eluitkak Pass, Elson Lagoon near — 


Point Barrow, 13 fms., gravel; off Point Barrow base, 21-78 fms., on 
bottoms of gravel, stones, small rocks, with shells and worm tubes (8 
stations, 12 specimens). Canap1an Arctic: Ellesmere Island, U.S.S. 
Alert, 1950, J. Peter Johnson (epitoke). East GREENLAND: 74°04’ N., 
17°58’ W., 120 fms., Norcross-Bartlett Expedition, 1931. Hast 
Coast Norts America: Off Labrador, 9-100 fms., mud, Blue Dolphin 
Expeditions, 1950, 1951. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Norway, 
Spitsbergen, Franz Josef Land, Barents Sea, Novaya Zemlya, Kara 
Sea. Also Iceland, Faroes; Hudson Bay to Labrador; Bering Sea to 
Oregon; north Japan Sea. In low water to 439 fathoms; epitokes at 
surface. Variety ? procera Fauvel (1914, 1923; not Ehlers, 1868): 
English Channel to Mediterranean. Variety persica Fauvel: Red 
Sea, South Arabian Coast, Persian Gulf, Indian Ocean. 


Family NEPHTYIDAE 


Body elongated, subtetragonal in cross section. Prostomium small, 
somewhat flattened, angulate, with four small antennae (fig. 30, 7). 
First segment rudimentary, with setae and usually with short dorsal 
and ventral tentacular cirri. Parapodia biramous, with rami well 
separated, with small dorsal cirri on lower sides of notopodia; with 


branchiae (interramal cirri) between the two rami, cirriform, sickle- 


shaped or foliaceous; with simple setae arranged in fan-shaped bundles; 


MARINE POLYCHAETE WORMS—PETTIBONE 267 


ventral cirri short, conical, or foliaceous; with anterior and posterior 
lamellae more or less developed (fig. 30, k-n). A single anal cirrus. 
Proboscis eversible, strong, muscular, a pair of horny jaws within the 
pharynx, with rows of soft papillae and a crown of terminal, bifid 
papillae. Move rapidly with undulating motion; usually in sand 
or mud. 

Represented by a single genus and four species. 


Genus Nephtys Cuvier, 1817 


The four species represented have numerous segments (65-150), 
tapered posteriorly and slightly anteriorly. Prostomium pentagonal 
to quadrate, with antennae conical, subequal, or more ventral pair 
slightly larger, without eyes, with pair of nuchal papillae at postectal 
margins (may be inverted or everted; fig. 30, 7). Tentacular or first 
segment with both dorsal and ventral tentacular cirri. Parapodial 
rami with anterior lamellae rudimentary or lacking, with posterior 
lamellae developed in varying degrees, with branchiae recurved (con- 
vex side toward lateral side of body, fig. 30, k-n). Ventral cirri short, 
conical. Proboscis with 22 long terminal papillae (10 pairs bifid, and 
a single dorsal and ventral one), with 22 longitudinal rows of sub- 
terminal papillae, four to eight per row, decreasing in size basally. 


Key to the species of Nephtys from Point Barrow 


1. Branchiae cirriform, sickle-shaped, with dorsal veirri digitiform (fig. 30, 1, n). 
Proboscis with a long median dorsal papilla._...----------------------- 2 
Branchiae foliaceous or cirriform and inflated basally, rudimentary or absent 
on last 15-30 segments, with dorsal cirri flattened, triangular, pointed (fig. 
30, k, m). Proboscis without a long median dorsal papilla; proximal surface 
of proboscis ridged (may be low tubercles in N. discors) ----------------- 3 
2. Branchiae begin on setigers 3 or 4, continuing to almost posterior end; acicular 
lobes rounded or slightly bilobed, especially anteriorly (fig. 30, l). Neuro- 
podial postsetal lamellae extend well beyond the acicular lobes, bilobed or 
irregularly sinuous. Setae very long, flowing. Proximal surface of pro- 
boscis smooth or furrowed_______+--.---=----==---------- N. longosetosa 
Branchiae begin on setigers 5-8, rudimentary on last 20-30 segments. Acicular 
lobes deeply bilobed in anterior and median regions (fig. 30, n). Neuropodial 
postsetal lamellae about same length as or only slightly surpassing the 
acicular lobes. Setae shorter. Proximal surface of proboscis tubercled. 
N. ciliata 
3. Branchiae begin on setiger 6, inflated basally, digitiform, recurved distally 
(fig. 30, m). Acicular lobes unequally bilobed, rounded in posterior region. 
Postsetal lamellae of anterior and median regions large, foliaceous, extending 
well beyond the acicular lobes; in posterior region, extend only slightly 
begend?:- te bene soe. ce eb teed - - accel sere sees - N. discors 
Branchiae begin on setigers 8-14, thick, wide, foliaceous (fig. 30, k). Acicular 
lobes rounded to conical. Postsetal lamellae shorter than, as long as, or 
only slightly longer than the acicular lobes_----------------- N. paradoxa 


261112—54——_5 


268 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Nephtys longosetosa Oersted, 1843 
Ficure 30, l 


Nephtys longosetosa Oersted, 1843, p. 195, pl. 6, figs. 75, 76.—Hartman, 1944a, 
p. 339, pl. 15, fig. 7. 

Nephthys longisetosa not Malmgren, 1865, p. 106, pl. 12, fig. 20—Eliason, 1920, 

. 32. 

Nephthys coeca Murdoch, 1885, p. 153 (part; includes var. longisetosa; not Nereis 
caeca Fabricius, 1780). 

Nephthys longosetosa Fauvel, 1923, p. 367, fig. 143, f-h—Monro, 1928, p. 82.— 
Annenkova, 1937, p. 163; 1938, p. 162.—Friedrich, 1939, p. 123.— Berkeley 
and Berkeley, 1943, p. 130; 1948, p. 52, fig. 76.—Gorbunov, 1946, p. 38.— 
Wesenberg-Lund, 1950a, p. 21; 1950b, p. 60; 1951, p. 44. 

Nephthys ciliata forma longosetosa Augener, 1928, p. 698. 

Nephthys ciliata Treadwell, 1937, p. 28 (part; not Nereis ciliata Miller, 1789). 

Description.—Length up to 170 mm., width without setae 6 mm. 
Tentacular cirri subequal to antennae. Notopodial postsetal lamellae 
unequally bilobed (a rather large upper lobe and small lobe just above 
the dorsal cirri), extending slightly or not beyond the acicular lobes. 
Neuropodial postsetal lamellae of anterior region rounded, entire, 
extending beyond the acicular lobes; those of the median and posterior 
regions large, bilobed or irregularly sinuous on ventral side, extending 
well beyond the acicular lobes. Cotor: In alcohol: Without color, 
or dusky, especially anteriorly. 

New records.—Arctic AuasKA: Eluitkak, Elson Lagoon near Point 
Barrow; Point Barrow base, washed ashore; off Point Barrow base, 
1.7-25 fms., on bottoms of gravel, stones, and mud (7 stations, 16 
specimens) ; Cape Lisburne, 5-7 fms., Dall. Brrine Sra: 60°16’ N., 
167°41’ W., Stoney, 1884; 66°45’ N., 166°35’ W., 10 fms., Dall, 1880. 
CanapiAn Arctic: Duckett Cove, Hurd Channel, Melville Peninsula, 
Foxe Channel, Bartlett, 1933. East Coast Norra America: Off 
Labrador, 8 fms., Blue Dolphin Expedition, 1949. 

Distribution — Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Barents Sea, White Sea, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes to France; to Maine; to Lower California and Panamé; north 
Japan Sea; Strait of Magellan. In low water to 528 fathoms. 


Ficure 30.—Nereidae: a, Nereis pelagica, dorsal view anterior end; 6, same, middle para- 
podium; c, Nereis zonata, middle parapodium; d, compound heterogomph falciger; e, 
compound homogomph falciger; f, compound heterogomph spiniger; g, compound homo- 
gomph spiniger; h, Nereis zonata, dorsal view extended proboscis showing areas of parag- 
naths; 4, same, ventral view. Nephtyidae: 7, Nephtys paradoxa, dorsal view anterior 
end; k, same, middle parapodium, anterior view; 1, Nephtys longosetosa, middle para- 
podium, anterior view; m, Nephtys discors, middle parapodium, anterior view; n, Nephtys 
ciliata, middle parapodium, anterior view. (For explanation of symbols, see p. 210.) 


MARINE POLYCHAETE WORMS—PETTIBONE 269 


Ficure 30.—For explanation see facing page. 


270 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Nephtys ciliata (Miller, 1789) 
FiaureE 30, n 

Nereis ciliata Miller, 1789, p. 14, pl. 89, figs. 1-4. 

Nephthys ciliata Malmgren, 1865, p. 104, pl. 12, fig. 17—Webster and Benedict, 
1887, p. 709.—Moore, 1903, p. 433.—Eliason, 1920, p. 31.—Fauvel, 1923, 
p. 371, fig. 145, a-b; 1933, p. 39.—Augener, 1928, p. 699.—Gustafson, 1936, 
p. 7—Treadwell, 1937, p. 28 (part).—Annenkova, 1937, p. 164; 1938, p. 
162.—Friedrich, 1939, p. 123.—Gorbunov, 1946, p. 38.—Thorson, 1946, p. 
69, figs. 32, 33.—Berkeley and Berkeley, 1948, p. 55, fig. 82.—Wesenberg- 
Lund, 1950a, p. 21; 1950b, p. 59; 1951, p. 45. 

Nephthys caeca Murdoch, 1885, p. 153 (part; includes var. ciliata).—Moore, 1911, 
p. 243 (not Nereis caeca Fabricius, 1780). 

Nepthys ciliata Chamberlin, 1920, p. 9. 

Nepthys hudsonica Chamberlin, 1920, p. 10, pl. 2, figs. 4-6. 

Nephtys ciliata Hartman, 1944a, pp. 335, 339, pl. 15, fig. 9 (as N. incisa, not fig. 
10 (=N. caeca)); 1950, p. 95. 

Description.—Length 100-300 mm., width without setae 9-11 mm. 
Tentacular cirri subequal to antennae. Acicular lobes deeply bilobed 
in anterior and median regions; rounded or slightly bilobed in posterior 
region. Postsetal lamellae of both notopodia and neuropodia rounded, 
entire, short, about same length as or only slightly surpassing the 
acicular lobes. Cotor: In alcohol: White, tannish, or anterior third 
reddish brown or buff. (According to G. E. MacGinitie, the pellets 
of this worm indicate that it is a detritus feeder.) 

New records—Arctic AuasKa: Off Point Barrow aero radio mast, 
10 fms.; off Point Barrow base, up to 8 miles from shore, 6-75.5 fms., 
on bottoms of mud and various combinations of mud, pebbles, gravel, 
stones, rocks, large perforated rocks, and shells (18 stations, 37 speci- 
mens). SrBERIA: Plover Bay, Dall, 1880. AtasxKa: Chiachi Island, 20 
fms., mud; Round Island, Coal Harbor, 6-8 fms., mud, 1872; Port 
Mulegrove, Yakutat Bay, 6-40 fms., 1874; Port Etches, 12-18 fms., 
1874; all collected by Dall. Wusr Coast Norra America: Wash- 
ington and Puget Sounds, low water to 83 fms. (common), Petti- 
bone. East Coast Nortu America: Off Labrador, 6-95 fms., Blue 
Dolphin Expeditions, 1949, 1950, 1951; off Nova Scotia, Bay of 
Fundy, Maine, Massachusetts, 14-110 fms., U. S. Fish Commission. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Norway, Barents Sea, 
White Sea, Novaya Zemlya, Kara Sea. Also Iceland, Faroes to 
France; to Massachusetts; to Southern California; to Japan, China. 
In low water to 500 fathoms. 


Nephtys discors Ehlers, 1868 
FiGureE 30, m 


Nephthys discors Ehlers, 1868, p. 626, pl. 23, figs. 39, 40.—Verrill, 1873, p. 103.— 
Webster and Benedict, 1887, p. 709. 

Nephtys discors Hartman, 1938c, p. 9, pl. 1, figs. 2, 3; 1950, p. 96. 

Nephtys rickettsti Hartman, 1938b, p. 153; 1950, p.'97. 


MARINE POLYCHAETE WORMS—PETTIBONE 271 


Description.—Length 115-300 mm., width without setae 9-12 mm. 
Dorsal pair tentacular cirri about half as large as ventral pair. 
Branchiae begin on setiger 6, at first a small swelling on the lower side 
of the base of the flattened, triangular dorsal cirrus, gradually becom- 
ing larger; where best developed, they are inflated on the basal half, 
with the distal half more slender, digitiform and slightly recurved or 
hooked; they are smaller and rudimentary only on posterior half of 
body. Notopodial acicular lobes of anterior and middle region 
unequally bilobed, the upper one wider and shorter and the ventral 
one narrower and longer; rounded in posterior region. Neuropodial 
acicular lobes slightly bilobed in anterior region; very low and rounded 
in middle and posterior regions (not very distinct from posterior 
lamellae). Conor: In alcohol: Brownish. 

Remarks.—The type of N. rickettsi Hartman from Alaska was 
compared with a specimen of N. discors from Bay of Fundy and is 
herein referred to the latter species. 

New records—Arctic ALAsKA: Point Barrow base, washed ashore; 
off Point Barrow base, up to 12.1 miles from shore, 24.7—123.5 fms., 
on bottoms of mud, gravel-mud, stones and large perforated rocks, 
and mass of worm tubes (5 stations, 5 specimens). Wrst Coast 
Nortu America: Puget and Washington Sounds, 70-83 fms., mud 
(3 specimens), Pettibone, 1938, 1939. East Coast NortH AMERICA: 
Bay of Fundy, U.S. Fish Commission. 

Distribution —Alaskan Arctic to southern California; Bay of Fundy 
to Maine. In 24-268 fathoms. 


Nephtys paradoxa Malm, 1874 
Ficure 30, j, k 

Nephthys paradoxa Malm, 1874, p. 78, pl. 1, fig. 2—Fauvel, 1923, p. 375, fig. 
146, f-i.—Augener, 1928, p. 701.—Gustafson, 1936, p. 7—Annenkova, 1938, 
p. 163.—Friedrich, 1939, p. 123.—Berkeley and Berkeley, 1943, p. 130.— 
Wesenberg-Lund, 1950a, p. 22; 1950b, p. 61; 1951, p. 47. 

Nephthys phyllobranchia McIntosh, 1885, p. 164, pl. 26, fig. 10; pl. 27, fig. 3; pl. 
14, A, figs. 12, 138. 

? Nephthys brachycephala Moore, 1903, p. 431.—Annenkova, 1937, p. 164; 1938, 
p. 162.—Zatsepin, 1948, p. 121, pl. 30, fig. 9. 

Nephtys paradoxa Hartman, 1944a, pp. 335, 339, pl. 15, fig. 6; 1950, p. 111. 

Nephtys phyllobranchia Hartman, 1950, p. 111. 

Description.—Length 150-200 mm., width without setae 8-13 mm. 
Dorsal pair tentacular cirri reduced to a tubercle, smaller than ventral 
pair. Branchiae begin on setigers 11-13 (8-14), very short, triangular 
at first, gradually becoming larger; where best developed, they are 
flat, wide foliaceous, weakly to distinctly concave; they are small, 
nearly absent on the last 15-30 segments. The anterior acicular 
lobes are rounded; those of the middle and posterior regions are 
conical, pointed. The postsetal lamellae in the anterior region are 


972 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


slightly longer than the acicular lobes; they are shorter than the 
conical acicular lobes in the middle and posterior regions. Cotor: 
In alcohol: Dusky, especially anteriorly. 

Remarks.—Augener (1928) indicated that he was of the opinion 
that N. phyllobranchia McIntosh should be referred to N. paradoza; 
Hartman (1950) stated that they are separable by only slight differ- 
ences. The type of N. brachycephala Moore from Japan is unsatis- 
factory (dried, when received); the description is in agreement with 
N. paradoxa. 

New records—Axrctic ALasKa: Off Point Barrow base, up to 12.1 
miles from shore, 23.5-123.5 fms., on bottoms of mud and mass of 
worm tubes (2 stations, 3 specimens). 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Norway, Barents Sea, 
Kara Sea. Also Iceland, Faroes to France; off Delaware; Japan. 
In 3.3-4,001 fathoms. 


Family GLYCERIDAE 


Body rounded, elongated, attenuated at both ends, not divided 
into two regions, either only uniramous or only biramous. Segments 
bi- or tri-annulate. Prostomium sharply conical, wide basally, with 
a pair of nuchal organs, transversely annulated, ending in four minute 
antennae distally (fig. 31, a). Small, globular dorsal cirri; larger 
ventral cirri (fig. 31, b, c); two anal cirri. Branchiae present or 
absent. Notosetae simple capillaries; neurosetae compound spinigers. 
Proboscis eversible, long, clavate, with numerous, small papillae 
(proboscidial organs) and four dark, falcate, horny jaws each with an 
attached, embedded, rodlike aileron (fig. 31, d). Epitoky affects 
entire individual (parapodial lobes become more elongate, setae longer 
and more numerous). 

Represented by a single species from Point Barrow. 


Genus Glycera Savigny, 1818 
Glycera capitata Oersted, 1843 
FiaureE 31, a-d 


Glycera capitata Oersted, 1843, p. 196, pl. 7, figs. 87, 88, 90-94, 96, 99.—Ehlers, 
1913, p. 503.—Fauvel, 1923, p. 385, fig. 151, a~-e—Augener, 1928, p. 734.— 
Monro, 1930, p. 115; 1936, p. 114.—Annenkova, 1937, p. 165; 1938, p. 164.— 
St¢gp-Bowitz, 1941, p. 186, fig. 1; 1948b, p. 4——Hartman, 1944a, pp. 336, 
339; 1948, p. 28; 1950, p. 76, pl. 11, figs. 1-4.—Gorbunov, 1946, p. 38.— 
Zatsepin, 1948, p. 117, pl. 31, fig. 9—Berkeley and Berkeley, 1948, p. 38, 
fig. 52.—Hartman and Reish, 1950, p. 20.—Wesenberg-Lund, 1950a, p. 23; 
1950b, p. 64; 1951, p. 49. 

Glycera setosa Oersted, 1843, p. 198, pl. 7, figs. 89, 95, 97 (epitokous form). 


MARINE POLYCHAETE WORMS—PETTIBONE PAB 


Glycera nana Johnson, 1901, p. 411, pl. 10, fig. 103.—Berkeley and Berkeley, 
1948, p. 37, figs. 50, 51. 

Glycera lapidum Fauvel, 1914, p. 205; 1923, p. 386, fig. 151, f-m; 1934a, p. 41.— 
Eliason, 1920, p. 36, fig. 6—Treadwell, 1937, p. 32.—Stgp-Bowitz, 1941, 
p. 191, fig. 2; 1948b, p. 7. 

Description.—Length 80 mm., width without setae 6 mm. (length 
up to 150 mm.—Hartman, 1950). Segments usually triannulate 
(may be biannulate, a slightly longer anterior ring at the level of the 
parapodia and a shorter posterior ring; the former may be sub- 
divided, thus triannulate; the latter may also be somewhat divided, 
thus tetra-annulate). Conical prostomium with 8-16 annuli (often 
difficult to count accurately). Dorsal cirri small, globular, inserted 
well above the parapodia. Ventral cirri wide, conical. Parapodia 
biramous throughout; notopodia represented by a short, conical 
presetal lobe and a small bundle of simple capillary setae; neuropodia 
with a longer, conical presetal lobe and a fan-shaped group of com- 
pound spinigerous setae; a single short, wide, rounded postsetal lobe 
common to both notopodia and neuropodia (thus two conical an- 
terior lobes and a single postsetal lobe). The relative length of the 
parapodial lobes may be variable, associated with the epitokous 
condition, as indicated by Hartman (1950). Proboscis with two 
kinds of papillae, numerous, long, cylindrical, slender ones, and 
fewer, larger, short, subspherical ones; aileron of jaws with a long 
lateral prolongation. Cotor: In alcohol: Brownish. 

Remarks.—Monro (1930) referred Glycera lapidum Quatrefages to 
G. capitata; they have been separated on minor and apparently 
variable characters; Fauvel (1914, 1923) regarded them as mere 
varieties. Hartman (1950) referred G. nana Johnson to G. capitata. 

New records—Arctic AtAska: Off Point Barrow base, 15 miles 
from shore, 78.2 fms., on bottom of small rocks with worm tubes (1 
station, 1 specimen). Norrawrest GREENLAND: Off Conical Rock, 
20-40 fms., 1938; 1 mile northwest Conical Rock, 1940; west side 
Wolstemholm Island, 1940; all collected by Bartlett. East Coast 
Norru America: Off Nova Scotia, Maine, Massachusetts. Rhode 
Island, low water to 410 fms., U. S. Fish Commission. ALASKA: 
Eastern Harbor, Sitka, gravelly-mud, 15 fms., 1874; between Pin- 
nacle and Ulakhta, Unalaska, 16 fms., 1874; Captain’s Harbor, 
Unalaska, 25-75 fms., coarse sand, 1874; all collected by Dall. 
Albatross Sta. Herendeen Bay, 1890. Wasuineton: Albatross Sta. 
2876, 48°33’ N., 124°53’ W., 59 fms., and Sta. 2879, 48°53’ N., 125°53’ 
W., 34 fms., 1888; Strait of Juan de Fuca, Washington and Puget 
Sounds, low water to 165 fms., Pettibone. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Davis Strait, Greenland, Spits- 
bergen, Norway, Barents Sea, White Sea, Novaya Zemlya, Kara 


274 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Sea. Also Iceland, Faroes to Portugal, Azores, Madeira, Mediter- 
ranean, Adriatic; to Rhode Island; to Gulf of California and Mexico; 
Japan; South Atlantic and Antarctic. In low water to 1,889 fathoms; 
epitokous phase in surface layers. 


Family GONIADIDAE 


Body elongated, more or less attenuated at both ends, divided 
into two regions, anterior region with uniramous parapodia (fig. 31, 
f) and long, wider posterior region with biramous parapodia (fig. 31, 
g)- Segments uniannulate. Prostomium sharply conical, trans- 
versely annulated, ending distally in four small antennae (fig. 31, e). 
Elongated conical dorsal and ventral cirri (fig. 31, f, g). Two anal 
cirri. Notosetae simple; neurosetae compound. Branchiae absent. 
Proboscis eversible, long, cylindrical, with terminal papillae and 
proboscidial organs, a pair of larger, dentate, dark horny macrognaths, 
and a circlet of many smaller H- or Y-shaped micrognaths. Epitoky 
involves only biramous portion. 

Represented by a single species from Point Barrow. 


Genus Glycinde F. Miiller, 1858 
Glycinde wiréni Arwidsson, 1898 
FicureE 31, e-g 


Goniada nordmanni Wirén, 1883, p. 403, pl. 30, figs. 4, 5, pl. 32, figs. 1, 2; (not 
Eone nordmanni Malmgren, 1867). 

Glycinde wiréni Arwidsson, 1898, p. 53, pl. 3, figs. 48, 49——not Moore, 1908, p. 
348 (=G. armigera Moore; in USNM). 

Description.—Length up to 45 mm., width without setae 2.5 mm., 
width including setae 5mm., 120 segments. Anterior region rounded, 
tapering sharply to pointed anterior end; posterior two-thirds wider, 
more flattened, tapering gradually posteriorly. Prostomium with 10 
annuli; two pairs of small deeply-set eyes, one pair in basal ring, one 
pair in distal ring (may be obscure). Anterior uniramous region 
consisting of 30-32 (31 in Arwidsson) setigers, with presetal lobe wide 
basally, narrowed abruptly to a lanceolate tip; postsetal lobe wide, 
conical; pre- and postsetal lobes as well as dorsal and ventral cirri 
subequal in length; a fan-shaped bundle of compound, spinigerous 
neurosetae. Posterior biramous region with shorter notopodia dis- 
tinctly separated from neuropodia. Notopodium a short, rounded 
lobe with a short dorsal cirrus and a short, protruding presetal lobe; 
notosetae few in number, acicular, hooked at tip, with long pointed 
hood. Neuropodia with short conical, postsetal lobe, subequal to 
ventral cirrus; presetal lobe wide basally with a lanceolate narrower 
tip, longer than postsetal lobe; neurosetae as in anterior region, 


MARINE POLYCHAETE WORMS—PETTIBONE 275 


except they may be longer and more numerous. Proboscis without 
chevrons; proboscidial organs (horny, yellow, spinous processes) of 
several kinds arranged in longitudinal series. Conor: In alcohol: 
Uniformly greenish brown or somewhat banded with one wide, dark- 
greenish-brown band per segment; iridescent. 

New records.—Arctic AtasKa: Off Point Barrow base, up to 4 
miles from shore, 23 to 29.1 fms., on various combinations of mud, 
sand, gravel, stones, rocks, shells (3 stations, 5 specimens). 

Distribution.—Arctic Alaska to Bering Sea. In 23 to 29 fathoms. 


Family LUMBRINERIDAE 


Body elongate, cylindrical, tapered slightly anteriorly and more 
strongly posteriorly, superficially resembling oligochaetes. Pro- 
stomium conical to rounded, with a pair of nuchal organs on posterior 
margin, without antennae or palps (fig. 31, h). First two segments 
apodous and achaetous, without tentacular cirri. Without dorsal 
and ventral cirri. Two to four anal cirri. Parapodia uniramous; 
Proboscis eversible, with dark, chitinous, well-developed masticatory 
apparatus consisting of a pair of ventral mandibles and four pairs of 
maxillary plates (fig. 31, 2, 7). 

Represented by a single species from Point Barrow. 


Genus Lumbrineris Blainville, 1828 
Lumbrineris fragilis (O. F. Miiller, 1776) 
Figure 31, h-n 


Lumbricus fragilis Miller, 1776, p. 216; 1788, p. 22, pl. 22, figs. 1-3, 

Lumbrinereis fragilis Malmgren, 1867, pl. 14, fig. 83.—Chamberlin, 1920, p. 15.— 
Eliason, 1920, p. 33.—Berkeley and Berkeley, 1943, p. 130. 

Lumbriconerets minuta Théel, 1879, p. 42, pl. 4, figs. 57-59.—Fauvel, 1911, p. 
22.—Augener, 1928, p. 732.—Wesenberg-Lund, 1950a, p. 28; 1950b, p. 71. 

Lumbriconerets fragilis Fauvel, 1923, p. 430, fig. 171, k, 1—Augener, 1928, p. 
730, pl. 11, fig. 5—Annenkova, 1937, p. 167; 1938, p. 167.—Gorbunov, 1946, 
p. 39.—Thorson, 1946, p. 74.—Zatsepin, 1948, p. 124, pl. 31, fig. 14— 
Wesenberg-Lund, 1950a, p. 27; 1950b, p. 69; 1951, p. 55. 

Lumbrineris fragilis Hartman, 1944a, pp. 335, 340. 


Deseription.—Length 150-250 mm., width without setae up to 7 
mm., width with setae 12 mm. Prostomium conical, pointed. First 
two achaetous segments about equal to the following segments or the 
first one slightly longer. Parapodia without branchiae, with presetal 
lobe short, rounded; postsetal lobe in anterior region of body slightly 
longer, rounded, diagonally truncate; in middle and posterior regions 
of body, postsetal lobe extended somewhat dorsally, somewhat digiti- 
form; with arched limbate setae with fine capillary tips, dark amber- 
colored on basal half, on first 60-100 setigers; with simple (not com- 
pound), hooded, hooked setae, dark amber-colored basally, beginning 


276 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


on setigers 15-24 (22-35 or more); on young specimens, the hooded, 
hooked setae may begin on more anterior segments; acicula black. 
With two anal cirri on ventral side of short, disclike pygidium. Jaw 
formula of proboscis (numbers refer to number of teeth): M II, 4+4 
(sometimes 5?); M III, 1+1 (or 2+1); MIV, 1+1. Cotor: In life: 
Reddish orange, iridescent. In alcohol: Colorless to gray, iridescent; 
young specimens uniformly reddish brown or somewhat banded with 
one wide dark brown band per segment. 

Remarks.—Three small specimens (0.8 mm. wide without setae) 
have the hooded, hooked setae beginning on the first one to three 
setigers as in L. minuta Théel. They are considered to be the young 
of L. fragilis as has been suggested earlier by Levinsen (1882) and 
Eliason (1920). 

New records—Arctic Atasxa: Off Point Barrow base, up to 12.1 
miles from shore, 20—123.5 fms., on bottoms of mud, stones, worm tubes 
and various combinations of mud, sand, gravel, stones, rocks, shells 
(16 stations, 29 specimens). East Gremntanp: Off Cape Hold with 
Hope, 23-40 fms., Bartlett, 1939. Eastern Norra America: Off 
Labrador, 5-13 fms., Blue Dolphin Expeditions, 1949, 1950; off Nova 
Scotia, Bay of Fundy, Maine, Massachusetts, Rhode Island, 7-452 
fms., U. S. Fish Commission; Woods Hole region, low water and 
dredged, Pettibone, 1950, 1951. AuasKxa: Chiachi Islands, 20 fms., 
mud, Dall, 1874. 

Distribution.—Widely distributed in the Arctic: Siberian and Alas- 
kan Arctic, Davis Strait, Greenland, Jan Mayen, Spitsbergen, Franz 
Josef Land, Barents Sea, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Norway to Azores, Madeira, Mediterranean; Hudson Bay to 
Rhode Island; Bering Sea; north Japan Sea. In low water to 1,883 


fathoms. 
? Larval Lumbrineris fragilis (O. F. Miiller) 


On two occasions nonpelagic larval polychaetes were found in a 
mucous mass off Point Barrow base: September 9, 1948, 21 fathoms, 
from breaking rock (the larvae described as rose-red); September 6, 
1949, 36 fathoms, attached to tunicate Boltenia echinata. Prostomium 


Ficure 31.—Glyceridae: a, Glycera capitata, dorsal view prostomium; b, same, parapodium 
from anterior region, posterior view; ¢, same, parapodium from middle region, anterior 
view; d, same, one of jaws. Goniadidae: e, Glycinde wiréni, dorsal view anterior end; 
f, same, uniramous parapodium from anterior region, anterior view; g, same, biramous 
parapodium from posterior region, anterior view. Lumbrineridae: h, Lumbrineris 
fragilis, dorsal view anterior end; 7, same, maxillary plates of jaw apparatus (one of each 
pair shown); 7, same, mandibles of jaw apparatus; k, same, parapodium from anterior 
region, posterior view; /, same, parapodium from posterior region, posterior view; m, 
same, limbate seta with capillary tips; , same, simple hooded hooked seta (crotchet). 
(For explanation of symbols see p. 210.) 


MARINE POLYCHAETE WORMS—PETTIBONE 277 


aSevtenw ay dere eT s 
OO tie a - ms 


i J P N M 


Ficurs 31.—For explanation see facing page. 


278 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


conical, without appendages. Two wide peristomial segments, the 
second with wide transverse ciliated band. Setigerous segments 8, 
gradually tapering posteriorly, each parapodium with single limbate 
capillary seta and hooded seta. Pygidium bulbous, with wide trans- 
verse ciliated band. 


Family ORBINIIDAE (ARICIIDAE) 


Body long, vermiform, with very numerous segments, divided into 
two weakly to sharply separable regions: (1) thoracic: More or less 
flattened dorsoventrally and enlarged; attenuated anteriorly; neuro- 
podia cushionlike, with several rows of setae; notopodia more or less 
cirriform, a bundle of crenulate capillary setae (fig. 32, b); (2) ab- 
dominal: Much longer, semicylindrical; neuropodia bilobed, project- 
ing dorsally (fig. 32, c). Branchiae medial to notopodia, simple, 
dorsal, erect, lanceolate or straplike, strongly ciliated, a pair on all 
segments except the anterior ones. Prostomium conical or globular, 
without appendages, with or without two small eyes, with two ciliated 
nuchal organs (fig. 32, a). First one or two segments apodous and 
achaetous. Usually without dorsal or ventral cirri. Setae simple, of 
varied form. Pygidium with two to four anal cirri. Proboscis 
eversible, unarmed, a voluminous, weakly- to much-branched, soft sac. 

Represented by a single species from Point Barrow. 


Genus Scoloplos Blainville, 1828 
Scoloplos armiger (O. F. Miiller, 1776) 
FIGURE 32, a-e 


Lumbricus armiger O. F. Miiller, 1776, p. 215; 1788, p. 22, pl. 22, figs. 4, 5. 
Aricia arctica Hansen, 1882, p. 34, pl. 5, figs. 20-26.—Murdoch, 1885, p. 154. 
Scoloplos armiger Fauvel, 1914, p. 224; 1927, p. 20, fig. 6, k-g.—Eliason, 1920, 
p. 39.—Augener, 1928, p. 742.—Monro, 1930, p. 145.—Annenkova, 1931, p. 
203; 1937, p. 169; 1938, p. 170.—Okuda, 1937a, p. 102, fig. 4.-Wesenberg- 
Lund, 1939b, p. 12; 1950a, p. 29; 1950b, p. 73; 1951, p. 58.—Berkeley and 
Berkeley, 1942, p. 195; 1952, p. 97, figs. 197-199.—-_ Hartman, 1944a, pp. 336, 
340, pl. 18, fig. 5 (not pl. 19, fig. 6 (Orbinia)).—Thorson, 1946, p. 78, fig. 
37.—Gorbunov, 1946, p. 39.—Zatsepin, 1948, p. 128, pl. 32, fig. 1. 
Haploscoloplos alaskensis Hartman, 1948, p. 30, fig. 8, a—c. 


Description.—Length up to 120 mm., width 2mm. Prostomium 
conical, pointed, with two deeply buried eyes (not visible when pre- 
served). First segment achaetous and apodous. Branchiae first 
appear on setigers 12 or 13 (9-17), very small at first, then triangular, 
then ligulate. Thoracic region consisting of about 17 setigers 
(12-20); notopodia with conical postsetal lobes and numerous, finely 
crenulate setae with capillary tips; neuropodia elongate, cushionlike, 
with a median postsetal conical lobe, with numerous rows of crenulate 


MARINE POLYCHAETE WORMS—PETTIBONE 279 


setae with capillary tips; some of these setae in the lower part of the 
neuropodial bundle may be worn down, with blunt tips resembling 
crotchets. These so-called crotchets may be more or less numerous 
on all thoracic segments or on some only, or may be lacking. Ab- 
dominal region having an exceedingly straggly appearance with the 
dorsally directed branchiae and parapodial lobes; notopodia with 
erect, digitiform postsetal lobes and capillary crenulate setae and 
sometimes also with forked setae; neuropodia with two conical, 
unequal lips, with capillary crenulate setae and an elongate thickened 
parapodial flange below the neuropodial lobe. On the last few thoracic 
and first few abdominal segments there may be one or two extra 
conical podal lobes, one at the level of the lower part of the neuropodial 
setae and one subpodal; they are in the transitional region of thoracic 
and abdominal regions and become part of the parapodial flanges of 
the abdominal segments. Without intermediate interramal cirri, 
but with lateral, ciliated organs between the rami. Pygidium with 
two filiform anal cirri. Proboscis soft, more or less lobed. Conor: 
In alcohol: Without color or yellowish brown, with darker areas on 
the elongate neuropodial flanges of the abdominal region and on the 
branchiae, 

Remarks —Haploscoloplos alaskensis Hartman from southwestern 
Alaska is herein referred to S. armiger. It differs from the typical 
S. armiger in lacking crotchets in the thoracic neuropodia. As 
indicated previously by Fauvel (1914) and others, this seems to be a 
variable character, perhaps associated with the substratum and age. 
The crotchets were not present on small specimens from Point Barrow, 
and they may be more numerous in some parapodia than in others 
and more abundant in the lower part of the neuropodia; they appear 
to be worn crenulate capillary setae. On certain substrata, pre- 
sumably, the setae may not be worn and thus the so-called crotchets 
are absent. 

New records.—Arctic Ataska: Eluitkak Pass, Elson Lagoon near 
Point Barrow; west side Elson Lagoon near entrance to small lagoon 
to the west, 1.2 fms.; off Point Barrow base, up to 12.1 miles from 
shore, 1.7—75.5 fms., on bottoms of mud, stones, and various com- 
binations of mud, gravel, stones, rocks, large perforated rocks (12 
stations, 19 specimens). Between Icy Cape and Cape Lisburne, 
15-20 fms., mud, sand, Dall, 1874. Easr Coast NortH America: 
Bay of Fundy, Maine, Massachusetts, 13-48 fms., U. S. Fish 
Commission. 

Distribution—Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait, Greenland, Jan Mayen, Spitsbergen, 
Franz Josef Land, White Sea, Novaya Zemlya, Kara Sea. Also 
Iceland, Faroes, Shetlands to France, Mediterranean; Labrador to 


280 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Massachusetts; southern Alaska to British Columbia; Japan; South 
Pacific (Chile); Antarctic; off South Africa. In low water to 1,100 
fathoms. 


Family SPIONIDAE 


Body vermiform, without distinct regions. Prostomium small, 
elongated, scarcely separated from an enlarged peristomium lateral to 
prostomium, with or without eyes (usually four), antennae usually 
lacking (may be a small occipital antenna), two very long, extensile, 
tentaculiform, longitudinally grooved, easily deciduous palps (fig. 
32, f, 2,1, r). Parapodia biramous, with dorsal and ventral postsetal 
lamellae more or less developed; dorsal and ventral cirri lacking 
(fig. 32, g, s). With ligulate or cirriform dorsal branchiae on upper 
part of parapodial bases on a certain number of segments: (rarely 
pinnate, rarely lacking). Setae simple capillaries and hooded 
crotchets (fig. 32, h, k, g). Pygidium terminated in an anal cup 
(fig. 32, p) or anal cirri. Proboscis short, scarcely or somewhat 
protrusible, may be somewhat lobulated. 

Represented by four genera and four species from Point Barrow. 
All genera have branchiae present on certain segments and prostomia 
without frontal horns. 


Key to the genera of Spionidae from Point Barrow 


1. Setigerous segment 5 strongly modified with special large amber-colored 
setae and without dorsal and ventral lamellae (fig. 32, m, n). Branchiae 
numerous, paired, dorsal, simple, straplike. Notopodia with capillary 
setae only; neuropodia with capillary setae and hooded bidentate crotchets 
beginning on setigers 7 or 8 (fig. 32, g). Pygidium ending in anal cup 
Hip AD ip)" Sd dn WL Wee hi Ee hed eee. Dy eee eee Polydora (p. 280) 

Setigerous segment 5 not modified___..£*..._/_2-L___ 2-22. See eee 2 

2. Branchiae on only few anterior segments, 3-11 pairs, often pinnate (fig. 
32, 7), beginning on setiger 2. Capillary setae and hooded crotchets in both 
noto- and neuropodia. Pygidium with anal cirri-_-_-_-_- Prionospio (p. 282) 

Branchiae more numerous, simple, straplike. Notopodia with capillary setae 
only; neuropodia with capillary setae and hooded crotchets___._-.------ 3 

3. Branchiae begin on setiger 1, continuing up to last setigers. Pygidium with 

CNY Wal ys Glad el ile Pull IE, yap ac ay Pn apt eh cine ig eS ees Spio (p. 284) 
Branchiae begin on setiger 2, continuing on large number of segments. Py- 
gidium with anal cup, may be somewhat lobulated__-_- Nerinides (p. 285) 


Genus Polydora Bosc, 1801 
Polydora caulleryi Mesnil, 1897 
Figure 32, l-q 


Polydora caulleryi Mesnil, 1897, p. 88, pl. 3, figs. 12-16.—Fauvel, 1927, p. 54, 
fig. 19, f—h. 
Polydora brachycephala Hartman, 1936c, p. 48, figs. 3-5; 1944b, p. 258. 


MARINE POLYCHAETE WORMS—PETTIBONE 281 


Description—Up to 50 mm. long, 2.2 mm. wide. Prostomium 
feebly notched anteriorly or distinctly bifid, prolonged posteriorly in 
a crest extending to setiger 4 (to 6), without distinct occipital antenna, 
with four eyes in trapezoidal arrangement (or six in young specimens) 
or eyes lacking. Palps usually missing in alcohol. A protrusible, 
somewhat lobulated, brownish proboscidial region. Branchiae begin 
on setigers 7 or 8, continuing on numerous segments, absent from 
more posterior segments; branchiae short (about length of capillary 
notosetae), straplike, continuous with dorsal lamellae. Modified seti- 
ger 5 with group of capillary notosetae, usually 5-7 (3-12) special 
amber-colored hooks, and group of capillary neurosetae; special hooks 
all similar, falcate, with pectinate top. With conspicuous, somewhat 
conical dorsal and ventral postsetal lamellae on setigers 1—4, lacking 
on modified setiger 5, becoming rather low and wide on more posterior 
segments. With capillary setae in both notopodia and neuropodia; 
beginning on setiger 7, neuropodial capillary setae mostly replaced 
by hooded bidentate crotchets. In more posterior setigers (last 20 
or so), notopodia conical, lacking branchiae or lamellae, with conical 
bundle of awl-shaped setae scarcely surpassing notopodial tip and 
small bundle of capillary setae; neuropodia with two or three crotchets 
and few capillary setae. Anal disc with complete middorsal notch, 
shorter midventral and midlateral notches, thus 4-lobed. Cotor: In 
alcohol: Without color or grayish. 

Remarks —The type of P. brachycephala Hartman from central 
California, was examined and is herein referred to P. caulleryi. 

New records—Arctic AtasKka: Off Point Barrow base, up to 8 
miles from shore, 16.7—75.5 fms., on bottoms of stones and various 
combinations of mud, gravel, stones, rocks, large perforated rocks, 
with bryozoans and hydroids (8 stations, 16 specimens). West Coast 
Norra America: Washington and Puget Sounds, low water to 50 
fms., Pettibone. 

Distribution.—Arctic Alaska to central California; Ireland, Great 
Britain to France. In low water to 75.5 fathoms. 


Polydora larva 


A larva of a Polydora was taken in plankton August 2, 1948. This 
was perhaps the larva of P. caulleryi Mesnil since it was the only 
species of Polydora found in the Point Barrow region. Prostomium 
rounded, with two palps, and with black chromatophores. Fifth 
setigerous segment modified, with short, thick, slightly curved acicular 
setae; rest of 21 setigerous segments with long larval capillary setae. 
Pygidium a rounded disc, notched middorsally, with black chromato- 
phores.. Black chromatophores arranged transversely on setigers 2-5, 
then in two longitudinal lines on each side of middorsal line, extending 
to anal cup. 


282 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Genus Prionospio Malmgren, 1867 
Prionospio malmgreni Claparéde, 1868 


FIGuRE 32, i-k 


Prionospio malmgreni Claparéde, 1868, p. 333, pl. 22, fig. 3 (fide Fauvel, 1927).— 
Ehlers, 1913, p. 511.—Fauvel, 1927, p. 61, fig. 21, a-e.—Annenkova, 1937, 
p. 171; 1938, p. 174—Hartman, 1948, p. 36.—Wesenberg-Lund, 1951, 
p. 70.—Berkeley and Berkeley, 1952, p. 29, figs. 54, 55. 

Spiophanes tenuis Verrill, 1879b, p. 176. 

Prionospio tenuis Verrill, 1882, p. 370.—Hartman, 1944a, pp. 336, 340, ? pl. 19, 
fig. 7; not 1945, p. 32 (=P. treadwelli Hartman, 1951). 


Description —Length up to 25 mm., width 1 mm. Prostomium 
dilated anteriorly, with anterior border rounded, narrowing posteriorly, 
extending in obtuse crest to setiger 2 (to 4). Four eyes, two smaller, 
two much larger (may be lacking). Two long palps, usually missing 
(in alcohol). A short, eversible, somewhat lobulated proboscidial 
region. Branchiae four pairs, on setigers 2-5, completely separated 
from dorsal lamellae, pairs 1 and 4 longer, pinnate, somewhat decid- 
uous, pairs 2 and 3 shorter, smooth, triangular. Dorsal postsetal 
lamellae of first setiger small, much larger and triangular on next 
four setigers, then diminishing in size, becoming reduced beginning 
on setigers 8 or 9; ventral postsetal lamellae oval, rounded. Hooded, 
multidentate crotchets beginning about setigers 12-16 in neuropodia 
and about setiger 40 in notopodia. Beginning on setiger 7, slightly 
elevated transverse folds between dorsal lamellae, continuing on next 
6-14 segments. Pygidium with longer, unpaired dorsal cirrus and 
two smaller lateral ones. Without color in alcohol. 

Remarks.—The types of Prionospio tenuis Verrill from Cape Cod 
Bay were examined and are herein referred to P. malmgreni. There 
are four pairs of branchiae, the first and fourth pairs pinnate, the sec- 
ond and third pairs shorter, smooth. 

New records —Arctic ALAsKA: Off Point Barrow base, 4 miles from 
shore, 29 fms., on bottom of gravel, small stones (1 station, 2 speci- 


Ficure 32.—Orbiniidae: a, Scoloplos armiger, dorsal view anterior end; 5, same, parapodium 
from thoracic region, posterior view; c, same, parapodium from abdominal region, anterior 
view; d, ¢, same, crenulate neurosetae from thoracic region. Spionidae: f, Spio filicornis, 
dorsal view anterior end (palps missing); g, same, parapodium from anterior region, 
anterior view; 4, same, hooded bidentate crotchet; 1, Prionospio malmgrent, dorsal view 
anterior end; j, same, branchiae on setigers 2-5 (one of each pair shown); k, same, hooded 
multidentate crotchet; 1, Polydora caulleryi, dorsal view anterior end (palps missing); 
m, same, parapodium of modified fifth setigerous segment; n, same, special hooks from 
setiger 5; 0, same, notopodium of posterior setiger, with conical bundle awl-shaped 
setae; p, same, anal disc of pygidium, lateral view; g, same, hooded bidentate crotchet, 
r, Nerinides sp., dorsal view anterior end; s, same, parapodium from anterior region. 
(For explanation of symbols, see p. 210.) 


= 


MARINE POLYCHAETE WORMS—PETTIBONE 283 


Q R 


Ficure 32.—For explanation see facing page. 


261112—54—6 


284 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


mens). West Coast Norta America: Puget Sound, Washington, 
low. water, mud, Pettibone. East Coast Norta America: Off 
Labrador, 40 fms., Blue Dolphin Expedition, 1949; off New Hamp- 
shire and Massachusetts, 25-48 fms., U. S. Fish Commission. 

Distribution —Arctic Alaska to Washington; north Japan Sea; 
Iceland, Norway to Mediterranean; Labrador to Massachusetts; 
South Africa. In low water to 250 fathoms; surface. 


Genus Spio Fabricius, 1785 
Spio filicornis (Miller, 1776) 
Figure 32, f-h 

Nereis filicornis Miller, 1776, p. 218. 

Spio filicornis Malmgren, 1867, p. 91, pl. 1, fig. 1.—HEHliason, 1920, p. 40, figs. 
7-9.—Fauvel, 1927, p. 43, fig. 15, a-g—Augener, 1928, p. 738, pl. 11, fig. 7.— 
Annenkova, 1937, p. 169; 1938, p. 173.—Hartman, 194la, p. 293; 1948, p. 
36.—Berkeley and Berkeley, 1943, p. 130.—Thorson, 1946, p. 93, fig. 46.— 
Zatsepin, 1948, p. 132, pl. 32, fig. 11—Wesenberg-Lund, 1950a, p. 30; 
1950b, p. 76; 1951, p. 71. 

Spio limicola Verrill, 1879, p. 176. 

Spio filicornis var. pacifica Berkeley and Berkeley, 1936, p. 475; 1952, p. 25, figs. 
47, 48. 

Deseription.—Up to 30 mm. long, 2 mm. wide. Prostomium bell- 
shaped to conical, widest anteriorly, rounded, tapered posteriorly to 
short crest extending on first two setigers. Four eyes in trapezoidal 
arrangement, anterior pair larger. Two palps missing (in alcohol). 
A short, eversible, somewhat lobulated proboscidial region. Branchiae 
begin on setiger 1, continuing to posterior end, simple, straplike. 
Dorsal postseta] lamellae of more anterior segments short, wide, with 
branchiae fused to upper sides; in more posterior segments, lamellae 
cirriform, separate from branchiae. Ventral postsetal lamellae 
rounded, not notched. Capillary setae only in notopodia. Neuropodia 
with hooded, bidentate crotchets beginning on setigers 9-18, about six 
in number (5-10). Pygidium with four cirri? Cotor: In alcohol: 
Yellowish white with dark brown pigment on peristomial region and 
along each side of anterior fourth of body. 

Remarks.—The types of Spio limicola Verrill from Cape Cod Bay 
were examined and are herein referred to S. filicornis. S. limicola 
differs from S. setosa Verrill, to which it has been referred (Hart- 
man, 1942, p. 63), in the number of neuropodial hooded crotchets, 
6-9 in S. lamicola and about 16 in VS. setosa. 

New records—Arctic ALASKA: Off Point Barrow base, up to 8 
miles from shore, 36-75.5 fms., on various combinations of gravel, 
rocks, stones, large perforated rocks (2 stations, 2 specimens). West 
Coast Norra America: Washington Sound, low water, Pettibone. 
East Coast Norta America: Off Massachusetts, 15-31 fms., U. S. 
Fish Commission. 


MARINE POLYCHAETE WORMS—PETTIBONE 285 


Distribution —Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait, Greenland, Spitsbergen, Novaya Zemlya. 
Also Iceland, Faroes, Norway to France; Hudson Bay to Massachu- 
setts; Bering Sea to central California; Japan. In low water to 211 
fathoms; surface. 


Genus Nerinides Mesnil, 1896 
Nerinides sp. 


FigurE 32, 7, s 


Two small specimens obtained from plankton 10 feet from shore, 
agreeing in some respects with the description of N. cantabra Rioja 
(Fauvel, 1927, p. 31). Length 16 mm., width 1 mm., segments 57. 
Prostomium rounded anteriorly, tapering posteriorly, with middorsal 
crest extending on first setiger, with four eyes, one pair larger, a 
raised area just posterior to eyes (median antenna?), with two large, 
thick palps (fig. 32, 7). Dorsal and ventral postsetal lamellae distinct, 
dorsal ones may be slightly undulate, ventral ones entire, not notched 
(fig. 32, s). Branchiae begin on setiger 2, continuing to near posterior 
end, fused to upper part of dorsal lamellae. Body with low transverse 
dorsal ridges between bases of branchiae. First setiger with both 
dorsal and ventral setae. Notopodia with only capillary setae, being 
extra long in posterior region. Neuropodia with capillary setae and 
hooded bidentate crotchets beginning around setiger 20. Pygidium 
with 4-lobed membranous appendage. Without color in alcohol. 


Family CriRRATULIDAE 


Body more or less cylindrical, attenuated at both ends, with 
numerous, short, compact, nearly similar segments. Prostomium 
small, usually without appendages, with or without eyes. Pair of large 
grooved spioniform palps (fig. 33, d) or with more or less numerous 
grooved tentacular filaments inserted on dorsal surface of an anterior 
segment (fig. 33, a). Branchiae simple, filamentous, long, con- 
tractile, inserted above notopodia on few to many segments (may 
be absent). Parapodia biramous, lobes scarcely projecting (fig. 33, 
b). Setae usually simple, some capillary, others acicular crotchets 
(fig. 33, c). Without dorsal, ventral, or anal cirri. Proboscis smooth, 
unarmed. May have epitokous pelagic form at time of reproduction. 
Usually in mud; some (Dodecaceria) in calcareous matrix. 

Represented by two genera and two species from Point Barrow. 
Both genera have prostomium conical, buccal segment and two fol- 
lowing segments achaetous, more or less fused to prostomium, with 
lateral filamentous branchiae beginning just anterior to first setigerous 
segment, continuing on large number of segments. 


286 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Key to the genera of Cirratulidae from Point Barrow 


1. With few to numerous, grooved tentacular filaments just anterior to first 
setigerous segment, subequal to branchiae (without 2 palps; fig. 33, a). 

Cirratulus (p. 286) 

With 2 large grooved, spioniform palps just anterior to first setigerous segment 

(without tentacular filaments; palps deciduous; may be missing in epitoke; 

Bes SSA) 22a he ee ey ee Chaetozone (p. 287) 


Genus Cirratulus Lamark, 1801 
Cirratulus cirratus (Miiller, 1776) 


Figure 33, a—-c 


Lumbricus cirratus Miiller, 1776, p. 215. 

Cirratulus cirratus Malmgren, 1867, p. 95.—Chamberlin, 1920, p. 20.—Eliason, 
1920, p. 57.—Fauvel, 1927, p. 94, fig. 33, a-g; 1933, p. 46; 1936, p. 72.— 
Augener, 1928, p. 750, pl. 11, fig. 9—Monro, 1930, p. 154; 1936, p. 161.— 
Annenkova, 1931, p. 203; 1932, p. 136; 1934, p. 322; 1937, p. 173; 1938, p. 
181.—Berkeley and Berkeley, 1942, p. 197; 1948, p. 180; 1952, p. 31, figs. 
58, 59.—Hartman, 1944a, pp. 334, 341; 1944b, p. 263; 1948, p. 37.—Gorbunov, 
1946, p. 39.—Zatsepin, 1948, p. 133, pl. 32, fig. 18.—Wesenberg-Lund, 
1950a, p. 33; 1950b, p. 81; 1951, p. 74.—Hartman and Reish, 1950, p. 34. 

Cirratulus robustus Johnson, 1901, p. 423, pl. 14, figs. 149, 150.—Rioja, 1941, 
p. 728.—Berkeley and Berkeley, 1942, p. 197. 

Cirratulus cingulatus Johnson, 1901, p. 422, pl. 14, figs. 145-148.—Rioja, 1941, 

. 729. 

Sévanitte spectabilis Berkeley and Berkeley, 1952, p. 32. 

Description—Length up to 300 mm., width 5 mm. Prostomium 
variable in shape: Short, broad, bluntly conical; almost square- 
shaped; or tapered gradually, then abruptly. Prostomium with 
usually four or five (2-8) eyes on each side in obliquely transverse 
row or may be united in almost solid arc. Buccal segment more or 
less fused with prostomium, two following achaetous segments more 
or less distinct. Tentacular filaments anterior to first setigerous 
segment, in two groups of 2-24 in obliquely transverse rows, long, 
filamentous, grooved longitudinally. Branchiae begin just anterior to 
first setigerous segment, lateral to group of tentacular filaments, 
continuing through greater part of body, anteriorly inserted close to 
upper part of notopodia, then gradually shifting more dorsally and 
then slightly more ventrally. Parapodia consisting of slightly pro- 
jecting tori common to both noto- and neuropodia, with capillary 
setae in both lobes, with additional acicular crotchets first appearing 
in neuropodia and more posteriorly in notopodia also. Anus sub- 
dorsal. Cotor: In life: Orange yellow with red branchiae; dark 
orange brown with orange-red branchiae. In alcohol: Yellowish 
brown or without color. | 

New records—Arctic AuasKa: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 5-6.6 fms.; off Point Barrow base, up to 8 miles from 


MARINE POLYCHAETE WORMS—PETTIBONE 287 


shore, 13.3-75.5 fms., on bottoms of stones, and various combinations 
of mud, pebbles, rock, stones, gravel, large perforated rocks, with 
bryozoans, embedded in mud at bases of barnacles and bryozoans as 
Eucratea loricata (19 stations, 183 specimens). Si1perra: Plover Bay, 
10-35 fms., Dall, 1880. Brritne Sra: Albatross station, Nikolski, 
Bering Island, 1892; St. George Island, Pribilofs, 2 miles off shore, 
40 fms., Hanna, 1913; Atka Island, Aleutians, L. M. Turner, 1879. 
West Coast Norts America: Strait of Juan de Fuca, Washington 
and Puget Sounds, low water, Pettibone. East Coast Norta AmMER- 
1ca: Off Labrador, 95 fms., Blue Dolphin Expedition, 1949; off Maine, 
Massachusetts, low water to 67 fms., U. S. Fish Commission. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Barents Sea, Novaya 
Zemlya. Also Iceland, Farces, Norway to France, Canary Islands; 
Hudson Bay to Massachusetts; Bering Sea to México; north Japan 
Sea to Japan, Manchuria. Southern latitudes: Falkland Islands, 
Magellan Straits, South Georgia, Kerguelen. In low water to 1,611 
fathoms. 

Genus Chaetozone Malmgren, 1867 


Chaetozone setosa Malmgren, 1867 
FicurE 33, d 


Chaetozone setosa Malmgren, 1867, p. 96, pl. 14, fig. 84.—Théel, 1879, p. 54, pl. 
4, figs. 49-51—Fauvel, 1914, p. 217, pl. 20, fig. 1; 1927, p. 101, fig. 35; 
1934a, p. 47.—Eliason, 1920, p. 57.—Augener, 1928, p. 750.—Gustafson, 
1936, p. 8.—Monro, 1937, p. 301.—Annenkova, 1937, p. 174; 1938, p. 182.— 
Berkeley and Berkeley, 1942, p. 197; 1952, p. 35, fig. 63.—Hartman, 1944a, 
pp. 334, 341.—Zatsepin, 1948, p. 133, pl. 32, fig. 16.—Wesenberg-Lund, 
1950a, p. 34; 1950b, p. 81; 1951, p. 75. 

Description.—Length up to 25 mm., width 2 mm. Prostomium 
conical, acutely pointed, without eyes. Buccal segment and two 
following achaetous segments more or less fused; in specimens from 
Point Barrow, buccal segment fused with prostomium, second and 
third segments fused dorsally. Paired palps large, long, grooved, 
inserted just anterior to first setigerous segment, deciduous, often 
missing on preserved specimens and missing when in epitokous phase. 
Branchial filaments begin just anterior to first setigerous segment, 
immediately posterior to palps, continuing on large number of anterior 
segments, lacking on posterior region, readily deciduous, inserted just 
posterior to upper part of parapodial tori. Parapodia are slightly 
projecting tori common to both noto- and neuropodia. In anterior 
region, both lobes with long, delicate capillary setae; more posteriorly, 
notopodia with still longer capillary setae, neuropodia with shorter, 
stouter capillary setae and short, unidentate acicular crotchets; more 
posteriorly, with acicular crotchets in both noto- and neuropodia. 


288 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


In epitokous phase, dorsal setae in anterior region much longer; in 
posterior region, parapodial tori more projecting, dorsal and ventral 
acicular crotchets forming semicircular, fan-shaped group, almost en- 
circling body. Pygidium with small, rounded to conical lobe, ventral 
and posterior to anus (like a cover for anal opening). Conor: In 
alcohol: Light to dark gray, brownish, or iridescent bluish black. 

New records—Arctic ALAsKa: Off Point Barrow base, up to 12.1 
miles from shore, 20-123.5 fms., on various combinations of mud, 
stones, gravel, rocks, large perforated rocks, mass of worm tubes, on 
surface of tunicates (12 stations, 29 specimens). East Coast NortH 
America: Off Labrador, 95-125 fms., Blue Dolphin Expedition, 1949; 
off Maine, New Hampshire, Massachusetts, 11-58 fms., U. S. Fish 
Commission. 

Distribution —Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Davis Strait, Greenland, Spitsbergen, Franz Josef 
Land, Barents Sea, Novaya Zemlya. Also Iceland, Faroes, Norway 
to Canary Islands, Mediterranean, Adriatic, Gulf of Aden; Labrador 
to Massachusetts; Bering Sea to British Columbia; north Japan Sea. 
Southern latitudes: Falkland Islands, Magellan Straits, Kerguelen. 
In low water to 1,333 fathoms; surface. 


Family FLABELLIGERIDAE (CHLORAEMIDAE) 


Body subcylindrical or subfusiform, with short similar segments 
covered with papillae, often with adherent sand grains, may be 
encased in thick mucous mantle. Prostomium and buccal segment 
fused, with two thick grooved palps, two groups of filiform cephalic 
branchiae, and more or less developed buccal tube around mouth, 
all of which may be partially or completely retracted within anterior 
setigers, with or without setae of anterior setigers directed anteriorly 
forming a cephalic cage. Parapodia biramous; notosetae simple, 
capillary, ringed; neurosetae simple or pseudocompound, capillary 
or hooked. Without dorsal, ventral, or anal cirri. Anus terminal. 
Usually in muddy sand. 

Represented by two genera and three species from Point Barrow. 


Key to the genera of Flabelligeridae from Point Barrow 


1. Body covered with long pedunculate papillae (fig. 33, g), usually encased in 
thick mucous mantle. Fused prostomium and buccal segment partially 
retractile within first setigers and more or less hidden by cephalic cage formed 
by setae of first setiger, the setae long, directed anteriorly, almost completely 
encircling segmeént lic. luie_-0los. aliod_ 24. fence Flabelligera (p. 289) 

Body covered with short papillae, usually encrusted with sand, not encased 
in mucous mantle. Fused prostomium and buccal segment completely 
retractile within anterior setigers, may be completely hidden; without 
cephalic cage formed by anterior setigers; when buccal segment, etc. are 
retracted, anterior end is trilobed (fig. 33, h).-------------- Brada (p. 290) 


MARINE POLYCHAETE WORMS—PETTIBONE 289 


Genus Flabelligera Sars, 1829 


Flabelligera affinis Sars, 1829 
Ficur5B 33, e-g 


Flabelligera affinis Sars, 1829, p. 31, pl. 3, fig. 16 (fide Malmgren, 1867).—Malmgren, 
1867, p. 83.—Chamberlin, 1920, p. 21.—Eliason, 1920, p. 62.—Fauvel, 1927, 
p. 113, fig. 40, a-f.—Augener, 1928, p. 768.—Monro, 1930, p. 160, fig. 63; 
1936, p. 164.—Annenkova, 1937, p. 174; 1938, p. 184.— Berkeley and Berkeley, 
1943, p. 130.—Hartman, 1944a, pp. 335, 341, pl. 33, figs. 12, 13 (as ?Brada 
setosa Verrill).—Gorbunov, 1946, p. 39.—Zatsepin, 1948, p. 134, pl. 33, fig. 
1.—St¢gp-Bowitz, 1948a, p. 8, fig. 1; 1948b, p. 30.—Wesenberg-Lund, 1950a, 
p. 35; 1950b, p. 82; 1951, p. 76. 
Flabelligera infundibularis Johnson, 1901, p. 417, pl. 12, figs. 124-127.—Hartman, 
1938c, p. 14; 1948, p. 40.—Berkeley and Berkeley, 1952, p. 7, figs. 1-4. 
Flabelligera infundibularum Moore, 1923, p. 223. 
Flabelligera infundibuliformis Hartman and Reish, 1950, p. 35. 
Description — Length up to 130 mm., width including mantle 12-15 
mm.,segments up to71. Body rounded, attenuated slightly anteriorly 
and more so posteriorly. Body covered with numerous papillae 
which are elongate, thin, flexible, usually somewhat coiled, with larger 
bulbous tips quite variable in shape. Body usually covered with 
thick, mucous, transparent mantle (may be missing in small speci- 
mens or specimens appearing in surface waters) which is penetrated 
by the pedunculate papillae. Fused prostomium and buccal segment 
retractile within anterior setiger, with pair of stout, grooved palps, 
two groups of numerous filiform branchiae, two pairs of eyes (almost 
coalesced), and somewhat protruding buccal siphon around ventral 
mouth opening. First setigerous segment with wide, funnellike 
depression into which buccal siphon, etc., can mostly be withdrawn; 
with circlet of elongated, ringed setae around margin and directed 
forward or spread funnellike, forming cephalic cage; with numerous 
elongate papillae having slender straight peduncles and fusiform tips. 
Notopodia conical, diagonally truncate lobes, with capillary ringed 
setae, with elongate papillae on posterior side that may be up to three- 
fourths length of notosetae (similar in shape to those around cephalic 
cage; different from those on body). Neuropodia conical, each with 
usually one (sometimes two) large, hooked, amber-colored pseudo- 
compound seta, ringed basally, hooked tips dark to black. When 
mucous mantle is present, setae almost completely embedded and the 
hooked tips of neurosetae and capillary tips of notosetae only ex- 
tending beyond mantle. Pygidium a fleshy, rounded rim; anus 
subdorsal. Cotor: In alcohol: Without color or slightly reddish 
brown. 
New records—Anrctic ALASKA: West side Elson Lagoon near en- 
trance to small lagoon to west, 1.2 fms.; Point Barrow base, washed 
ashore; off Point Barrow base, 18.3-123.5 fms., on various combina- 


290 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


tions of mud, stones, gravel, rocks, with mass of worm tubes (12 
stations, 15 specimens). Brrine Sua: Albatross Sta. 3274, 55°34’ 
N., 162°31’ W., 19 fms., and station at Unalaska, Aleutians, 1890; 
Tluiliuk Harbor, Unalaska, Dall, 1871. Wrst Coast NortH AMERICA: 
Washington and Puget Sounds, 10 fms. and surface, Pettibone. 
CanapiaAn Arctic: Dobbin Bay, east Ellesmere Land, 79°36’ N., 
73°35’ W., 16 fms., W. H. Littlewood, 1950. Foxe Basin, Baffin 
Island, 66°83’ N., 80°07’ W., Bartlett, 1927. Wzxst GREENLAND: 
Between Capes Alexander and Chalon, 25-40 fms., Bartlett, 1937. 
East Coast Norra America: Off Labrador, 10-30 fms., Blue 
Dolphin Expeditions, 1949, 1950; Grand Manan, Maine, New 
Hampshire, Massachusetts, 3-90 fms., U. S. Fish Commission. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Barents Sea, White Sea, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Norway to France, Mediterranean; Hudson Bay to Mas- 
sachusetts; Bering Sea to southern California; north Japan Sea. 
Southern latitudes: Falkland Islands, Magellan Straits, South 
Georgia, South Africa, Australia. In low water to 889 fathoms; 
surface. 

Genus Brada Stimpson, 1854 


Key to the species of Brada from Point Barrow 


1. Body covered with filiform papillae, often encrusted with sand grains, with 
tips of papillae bare. Notopodia distinct, with 2-5 capillary setae per lobe. 

B. villosa 

Body covered with low globular papillae, often encrusted with a thin layer of 
sand grains. Notopodia indistinct, with 0-2 very delicate capillary setae. 

B. inhabilis 


Brada villosa (Rathke, 1843) 


Siphonostoma villosum Rathke, 1843, p. 215, pl. 11, figs. 11, 12. 

Brada villosa Malmgren, 1867, p. 84.—Eliason, 1920, p. 62.—Chamberlin, 1920, 
p. 22.—Fauvel, 1927, p. 121, fig. 43, e-1; 1934a, p. 49.—Monro, 1930, p. 161; 
1937, p. 303.—Annenkova, 1937, p. 175; 1938, p. 185.—Okuda, 1937b, 
p. 53, pl. 2, fig. D.—Berkeley and Berkeley, 1943, p. 130; 1952, p. 7, fig. 5.— 
Zatsepin, 1948, p. 136, pl. 33, fig. 4—Stgp-Bowitz, 1948a, p. 33, fig. 9; 
1948b, p. 39, fig. 15.—Hartman and Reish, 1950, p. 35.—Wesenberg-Lund, 
1950a, p. 35; 1950b, p. 85; 1951, p. 78, fig. 7. 

Brada setosa Verrill, 1873, pp. 431, 434, 508, 606; 1881, p. 302, pl. 9, fig. 4.— 
Sumner, Osburn and Cole, 1913, p. 630. 

Trophonia rugosa Hansen, 1882, p. 38, pl. 7, figs. 9-12. 

Trophonia arctica Hansen, 1882, p. 39, pl. 7, figs. 17-20. 

Brada granulata Murdoch, 1885, p. 155 (not B. granulata Malmgren, 1867). 

Brada pilosa Moore, 1906b, p. 231, pl. 10, figs. 14-17. 

Stylaroides pluribranchiata Moore, 1923, p. 222. 

Brada rugosa St¢p-Bowitz, 1948a, p. 37, fig. 10; 1948b, p. 41. 


MARINE POLYCHAETE WORMS—PETTIBONE 291 


Description.—Length 28-60 mm., width 5-9 mm., setigers 23-35. 
Body fusiform, slightly flattened ventrally, convex dorsally, attenu- 
ated and more or less elongated posteriorly, having a characteristic 
rough, hirsute aspect, covered with cylindrical, fusiform to claviform 
papillae, longer dorsally than ventrally, usually encrusted with sand 
grains more basally, with distal tips bare; there are all gradations: 
Without encrusting sand (looks “‘hairy”’), with scattered sand grains, 
and with rather thick layer of sand grains, usually fewer on antero- 
ventral part. Underlying surface nodular and, in larger specimens, 
outer papillate layer may be worn off, the surface appearing rugose, 
with low mounds. Retractile prostomium and buccal region with two 
groups of numerous filiform branchiae borne on pair of dorsolateral 
bosses, pair of thick grooved palps, and ventral eversible buccal siphon 
around mouth. Setae of first setigerous segment 8-10 per bundle, 
slightly more elongate than following, directed anteriorly. Notopodia 
and neuropodia distinct, with long fusiform papillae grouped around 
setae. Notosetae long, capillary, ringed, 2-5 in number. Neurosetae 
beginning on setiger 2, large, 3-6 in number, amber-colored, ending in 
sharp, transparent, and fragile tips. Pair of nephridial papillae near 
anterior border of ventral side of setiger 5. Conor: In life and in 
alcohol: Brownish, dull sandy-mud. 

Remarks.—The types of Brada setosa Verrill from Massachusetts, 
B. pilosa Moore from Alaska, and Stylaroides pluribranchiata Moore 
from California were examined and are herein referred to B. villosa. 
The excessive “hairiness” of B. pilosa appears to be a variable charac- 
ter; there may be varying numbers of papillae as well as all gradations 
in the amount of encrusting sand grains. Two large specimens washed 
ashore at Point Barrow were at first identified as B. rugosa, following 
Stgp-Bowitz (1948) ; they agree in the large size (23-24 setigers, 28-30 
mm. long, 5-7 mm. wide), the almost complete absence of papillation, 
and the thick rugose surface; the outer surface, however, appears to be 
worn off. There are some remains of the papillated surface on the 
parapodial lobes. Thus B. rugosa (including TJ. arctica) is herein 
referred to B. villosa, where it has been referred earlier by Fauvel and 
others. 

New records—Arctic ALasKa: Point Barrow base, washed ashore 
September 22, 1949 (22 specimens), August 10, 1950 (2 specimens) ; off 
Point Barrow base, up to 4 miles from shore, 27-29 fms., on various 
combinations of mud, gravel, stones, rocks (2 stations, 2 specimens). 
Berrine Sua: 62°15’ N., 167°48’ W., Stoney, 1884; Albatross Sta. 3337, 
53°55’ N., 163°26’ W., 280 fms., 1890. Wexsr Coast Nortu AMERICA: 
Washington Sound, Pettibone; Albatross Sta., northwest West Point, 
Elliot Bay, Seattle, 1914. East Coasr Norra America: Off Nova 


292 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Scotia, Maine, Massachusetts, Rhode Island, 6-499 fms., U. S. Fish 
Commission. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Barents Sea, Novaya 
Zemlya, Kara Sea. Also Iceland, Faroes, Norway to Spain, Mediter- 
ranean, Adriatic, south Arabian Coast; Hudson Bay to Rhode Island; 
Bering Sea to southern California; north Japan Sea to Japan; South 
Atlantic: South Orkney and South Shetland Islands. In low water 
to 853 fathoms. 


Brada inhabilis (Rathke, 1843) 
FicurE 33, h 


Siphonostoma inhabile Rathke, 1843, p. 218, pl. 11, fig. 13. 

Brada sublaevis Stimpson, 1854, p. 32.—Verrill, 1881, pp. 289, 304.—Webster 
and Benedict, 1887, p. 731. 

Brada granulata Malmgren, 1867, p. 85, pl. 12, fig. 71.—Théel, 1879, p. 52.— 
not Murdoch, 1885, p. 155 (=B. villosa) —Annenkova, 1937, p. 176; 1938, 
p. 186.—Hartman, 1944a, pp. 334, 341; 1948, p. 41.—Zatsepin, 1948, p. 
135, pl. 33, fig. 3—Wesenberg-Lund, 1950a, p. 36; 1950b, p. 85. 

Brada granosa Treadwell, 1937, p. 32 (not B. granosa Stimpson, 1854). 

Brada inhabilis St¢ép-Bowitz, 1948a, p. 40, fig. 11; 1948b, p. 42, fig. 16.—Wesen- 
berg-Lund, 1951, p. 80. 


Description.—Length 30-60 mm., width 8-12 mm., segments 23-26. 
Body subfusiform, grub-shaped, flattened ventrally, strongly arched 
dorsally, usually curved ventrally (in alcohol). Body surface covered 
with low globular papillae (to naked eye surface appears almost 
smooth, granular), usually with thin layer of sand grains, often with 
an almost uniform layer of larger sand grainson dorsum. Prostomium 
and buccal segment with two groups of three to four filiform branchiae, 
pair of grooved palps, and buccal siphon around mouth. First seg- 
ment achaetous (may be few very, delicate capillary setae), second 
segment with two small neurosetae, rest of segments with neuropodia 
as slightly elevated lobes and about four (2-9) amber-colored neuro- 
setae with curved tips. Without distinct notopodia; notosetae poorly 
developed, one or two or absent, extremely capillary, easily overlooked. 
Nephridial papillae on ventral side between segments 4 and 5 (setigers 3 
and 4, since the first segment is usually achaetous). Cotor: In life: 
Orange yellow. In alcohol: Light gray to ashy. 

Remarks.—B. granulata Malmgren is herein referred to B. inhabtlis, 
following Stgp-Bowitz (1948a). B. sublaevis Stimpson should also be 
referred here and not to B. rugosa (Hansen) as indicated by Stgp- 
Bowitz (1948a, p. 37). 

New records —Anrctic ALASKA: Off Point Barrow base, up to 12.1 
miles from shore, 20—-123.5 fms., on bottoms of rocks, stones, gravel, 
and mass of worm tubes (5 stations, 9 specimens). NoRTHWEST 


MARINE POLYCHAETE WORMS—PETTIBONE 293 


GREENLAND: 1 mile northwest Conical Rock, 25-60 fms., Bartlett, 
1988. Brrine Sea: Albatross Sta. 3223, 54°26’ N., 165°32’ W., 
56 fms., 1890; Bering Strait, 13 fms., Dall, 1880. Azasxa: Iliuliuk 
Harbor, Unalaska, 1871, and New Harbor, Unga Island, under stones, 
1872, Dall; Kodiak, W. J. Fisher. East Coast Norra America: 
Off Labrador, 95 fms., Blue Dolphin Expedition, 1949; Bay of Fundy, 
Maine, 48-90 fms., U. S. Fish Commission. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Novaya Zemlya,Kara 
Sea. Also Iceland, Faroes, Norway to Danish waters; Labrador to 
Maine; Bering Sea to Gulf of Alaska; north Japan Sea. In low water 
to 609 fathoms. 


Family SCALIBREGMIDAE 


Body inflated anteriorly or short, fusiform. Integument roughened 
or tesselated and divided into superficial rings. Prostomium small, 
bilobed, or entire and with frontal horns; two ciliated evaginable 
nuchal grooves on each side of prostomium (fig. 33, 7). Buccal 
segment achaetous. Parapodia biramous, rami small, with or without 
dorsal and ventral cirri. Setae simple, capillary, some single, some 
forked. Branchiae limited to anterior few segments (may be absent). 
Pygidium with or without anal cirri. Proboscis eversible, unarmed, 
globular, smooth. 

Represented by a single species from Point Barrow. 


Genus Scalibregma Rathke, 1843 
Scalibregma inflatum Rathke, 1843 
Figure 33, i-k 


Scalibregma inflatum Rathke, 1843, p. 184, pl. 9, figs. 15-21.—Moore, 1923, 
p. 217.—Fauvel, 1927, p. 123, fig. 44,a-f; 1932, p. 187; 1934a, p. 50.— 
Augener, 1928, p. 754.—Monro, 1930, p. 163.—Annenkova, 1934, p. 322; 
1937, p. 176; 1938, p. 186.—Okuda, 1938b, p. 99.—Hartman, 1944a, pp. 
336, 341; 1948, p. 40.—Stgp-Bowitz, 1946, p. 67, fig. 2; 1948b, p. 24, fig. 8.— 
Zatsepin, 1948, p. 136, pl. 33, fig. 10—Wesenberg-Lund, 1950a, p. 36; 
1950b, p. 86; 1951, p. 81.—Berkeley and Berkeley, 1952, p. 58, figs. 119-121. 

Scalibregma brevicauda Verrill, 1873, pp. 416, 422, 605; 1881, p. 302, pl. 9, fig. 5. 

Scalibregma minutum Webster and Benedict, 1887, p. 727. 


Description.—Length up to 100 mm., width 13 mm., 45-60 segments 
(two specimens washed ashore after a storm were longer than any 
previously recorded). Body divided into two more or less distinct 
regions, anterior inflated portion (anterior 15-17 setigers) tapered 
anteriorly, without dorsal and ventral cirri, and narrower attenuated 
posterior region with dorsal and ventral cirri. A more or less devel- 


294. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


oped ventral groove along length of body, better developed in posterior 
region. Integument tesselated, with four annuli per segment. Pro- 
stomium subquadrangular, wider anteriorly, with frontal horns on 
anterolateral borders, with pair of evaginable nuchal organs lateral 
to prostomium (when inverted, appear as diagonal slits). Buccal 
segment achaetous, forming rugose ring around prostomium. Mouth 
ventral, at level of first setigerouslobes. Proboscis eversible, globular, 
smooth, unarmed. Branchiae four pairs, short, tufted, arborescent, 
inserted posterior to notopodia on setigers 2—5, increasing in size 
posteriorly. Noto- and neuropodial lobes of anterior segments small, 
projecting, subequal. Beginning on setigers 16-18, dorsal and ventral 
cirri develop gradually, becoming elongate-conical, flattened or 
inflated. Setae silky, iridescent, capillary (may be extra long in 
epitokous pelagic form). Lateral ciliated organs between rami. 
Pygidium with five'to seven, short to long, deciduous, filiform anal 
cirri on ventral side. Epitokous sexual forms may appear in surface 
waters, massed with sexual products, with extra long setae and 
usually with highly pigmented dorsal and ventral cirri. Cotor: 
In life: Orange yellow. In alcohol: Without color, orange yellow, or 
sulfur yellow with darker pigmented masses in the outer portions of 
the dorsal and ventral cirri. 

Remarks.—Scalibregma brevicauda Verrill is herein referred to S. 
inflatum. Although the types were not examined, specimens from 
the New England region identified by Verrill as S. brevicauda were 
observed. The types of S. minutum Webster and Benedict from 
Eastport, Maine, were examined and found to agree with S. inflatum. 

New records—Arctic ALASKA: Point Barrow base, washed ashore; 
off Point Barrow base, 22.5-78 fms., up to 15 miles from shore, on 
bottoms of mud and various combinations of mud, sand, gravel, 
stones, rocks, with shells and worm tubes (9 stations, 17 specimens). 
West Coast Nortru America: Washington and Puget Sounds, 25 
fms., mud, Pettibone. East Coast Norra America: Off Labrador, 
17-35 fms., Blue Dolphin Expeditions, 1950, 1951; off Nova Scotia, 
Maine, Massachusetts, Rhode Island, Connecticut, Long Island 
Sound, 5-95.5 fms., U. S. Fish Commission; Cuttyhunk, Massachu- 
setts, low water, in sand, Pettibone, 1950. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Jan Mayen, Spits- 
bergen, Franz Josef Land, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Norway to France, Adriatic, Gulf of Oman; Labrador to 
Long Island Sound; Bering Sea to central California; north Japan 
Sea to Japan. Southern latitudes: New Zealand, Magellan Straits, 
South Georgia, Prince Edward Islands, Kerguelen. In low water to 
1,333 fathoms; epitokes at surface. 


MARINE POLYCHAETE WORMS—PETTIBONE 295 


Family OPHELIIDAE 


Body relatively short, often grublike or lancet-shaped. Integument 
smooth or granulated; segmental grooves weak or obscured by super- 
ficial annuli. Prostomium small, conical, without appendages; pair 
of eversible nuchal organs (fig. 33, 1, 0). Proboscis eversible, saclike, 
unarmed. Parapodia biramous, lobes reduced or lacking; setae simple, 
capillary (fig. 33, m, q). Without dorsal cirri; rarely with ventral 
cirri. Branchiae paired, cirriform, inserted near parapodia, extending 
along most of body (may be lacking). Pygidium forming an anal 
cylinder, a flap, or knob, with or without cirri (fig. 33, n, p). 

Represented by two genera and two species from Point Barrow. 
Both genera have paired cirriform branchiae along most of length of 
body; without lateral eyes along body. 


Key to the genera of Opheliidae from Point Barrow 


1. Body lancet-shaped; ventral surface thick, solelike, formed by 2 thick longi- 
tudinal muscular columns, with midventral and lateral grooves along length 

of body. Parapodial lobes small, along lateral grooves (fig. 33, m). 
Ammotrypane (p. 295) 
Body cylindrical, grub-shaped, without midventral or lateral grooves. Para- 
podial lobes absent, the simple, capillary noto- and neurosetae emerging 
directly from body wall (fig. 33, g)-------------------- Travisia (p. 296) 


Genus Ammotrypane Rathke, 1843 


Ammotrypane breviata Ehlers, 1913 
FIGuRB 33, l-n 


Ammotrypane breviata Ehlers, 1913, p. 523, pl. 39, figs. 1-7.—Monro, 1930, p. 
165; 1936, p. 165. 
Ophelina groenlandica St¢p-Bowitz, 1948b, p. 20, fig. 6. 

Description.—Length 6-7 mm., width 0.3 mm., setigerous segments 
26 or 27 (up to 34 mm. long, 2 mm. wide, 26-28 segments—Monro, 
1930, 1936). Body rigid, tapered at both ends, smooth, without 
superficial segmental divisions. Prostomium conical, the anterior tip 
constricted to form a small globular palpode; prostomial tip may be 
partly telescoped into rest of prostomium; pair of eversible nuchal 
organs at level of parapodia. Parapodia with two groups of fine 
capillary setae, the dorsal group slightly longer than the ventral. 
Branchiae begin on setiger 2, absent on last three to five setigers; 
branchiae slender, cirriform, subequal. Pygidium forming a short, 
somewhat contractile anal cylinder, closed on all sides, somewhat 
lobulate or papillate on open posterior end. LEversible proboscis 
saclike, smooth. Conor: In alcohol: Without color or may be darkly 
pigmented laterally along posterior three setigerous segments and 
proximal part of anal tube. 


296 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Remarks.—The two small specimens from Point Barrow (6-7 mm. 
long) follow closely the description of St¢gp-Bowitz for Ophelina groen- 
landica, including size. They seem to agree with the description by 
Ehlers of A. breviata from the Antarctic, although the Antarctic 
specimens attain a much larger size (29 mm. long). 

New records—Arctic AuasKa: Off Point Barrow base, up to 7.5 
miles from shore, 29-36 fms., on bottoms of gravel, stones, large 
perforated rocks (2 stations, 2 specimens). 

Distribution.—Arctic Alaska, East Greenland. Southern latitudes: 
South Orkney and Falkland Islands, Kaiser-Wilhelm II Land. In 
10-214 fathoms. 


Genus Travisia Johnston, 1840 


Travisia carnea Verrill, 1873 
FIGURE 33, 0-g 


Travisia carnea Verrill, 1873, pp. 431, 434, 508, 604; 1881, p. 302, pl. 8, fig. 1.— 
Sumner, Osburn, and Cole, 1913, p. 629.—Hartman, 1944a, pp. 336, 341, pl. 
19, fig. 5 (as probably Ophelia limacina). 
Travisia forbesi Murdoch, 1885, p. 154 (not 7. forbesit Johnston, 1840). 
Description.—Length up to 59 mm., width 8 mm., segments 25-29 
(specimens washed ashore at Point Barrow much larger than any 
previously recorded). Body rounded, cylindrical, grub-shaped, 
tapered toward each end. Segments of posterior third may be some- 
what imbricated or telescoped; triannulate in middle region. Integu- 
ment with granulations or beading rather uniform (not larger on 
posterior part of segment as in JT. pupa Moore). Prostomium small, 
conical, acutely to bluntly pointed, without eyes. Branchiae begin on 
setiger 2 (rarely on setiger 3), continuing posteriorly, absent on last 
four or five segments, cirriform, inserted posterior to notosetae. 
Without parapodial lobes, the small, slender, capillary noto- and neuro- 
setae emerging directly from body wall. In posterior third of body, 
area in region of setae may be somewhat inflated but without enlarged 
lateral lobes extending posteriorly (as in JT. forbesxi Johnston and 


Ficure 33.—Cirratulidae: a, Cirratulus cirratus, dorsal view anterior end; b, same, para- 
podium from anterior region; c, same, acicular crotchet; d, Chaetozone setosa, dorsal 
view anterior end. Flabelligeridae: ¢, Flabelligera affinis, parapodium; f, same, hooked 
pseudocompound neuroseta; g, same, pedunculate papillae—(1) from tip of notopodium, 
(2) on body; h, Brada inhabilis, ventral view anterior end. Scalibregmidae: i, Scalibregma 
inflatum, dorsal view anterior end; 7, same, parapodium from middle region; k, same, 
parapodium from posterior region. Opheliidae: 1, Ammotrypane breviata, ventral view 
anterior end, proboscis extended; m, same, lateral view middle part of body; , same, 
ventral view posterior end; 0, Travisia carnea, dorsal view anterior end; p, same, dorsal 
view posterior end; g, same, parapodium. Capitellidae: r, Capitella capitata, dorsal view 
anterior end; s, same, dorsal view posterior part thoracic region; ¢, same, parapodium 
from abdominal region; u, same, hooded crotchet. (For explanation of symbols, see 
p. 210.) 


MARINE POLYCHAETE WORMS—PETTIBONE 297 


Ficure 33.—For explanation see facing page. 


298 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


T. brevis Moore). With lateral sensory pits between rami appearing 
as small oval slits. Nephridial pores ventral to neuropodia on setigers 
3-14. Pygidium small, cylindrical, longitudinally furrowed. Cotor: 
In life: Flesh. In alcohol: Whitish. 

New records.—Axrctic AtasKa: Point Barrow base, washed ashore, 
October 16, 17, 1949; August 10, 1950 (12 specimens). East Coast 
Norta America: Off Massachusetts, Long Island Sound, 3-19 fms., 
soft mud, U.S. Fish Commission; Woods Hole region, Massachusetts, 
Pettibone, 1950. 

Distribution.—Arctic Alaska. Massachusetts to Long Island Sound. 
In 3-19 fathoms. 


Family CAPITELLIDAE 


Body cylindrical, weakly divided into 2 regions: Anterior thoracic 
region somewhat inflated, often reticulated, parapodial lobes rudi- 
mentary; posterior abdominal region longer and thinner, with rows 
of crotchets on somewhat inflated tori. Prostomium conical, more 
or less retractile, without appendages, with or without eyes, with 
pair of eversible nuchal organs. Buccal segment achaetous (may be 
fused with prostomium as in Capitella). Parapodia biramous, re- 
duced to bundles of capillary setae or to dorsal and ventral tori 
bearing row of crotchets which have long manubrium and recurved 
hooded rostrum (fig. 33, 7-w). Without dorsal, ventral, or anal cirri. 
With or without branchiae. Proboscis eversible, globular, unarmed, 
papillate. Constructs tubes. 

Represented by a single species from Point Barrow. 


Genus Capitella Blainville, 1828 
Capitella capitata (Fabricius, 1780) 
FIGURE 33, r—u 


Lumbricus capitatus Fabricius, 1780, p. 279. 

Capitella capitata Webster and Benedict, 1884, p. 730; 1887, p. 744.—Ehlers, 
1913, p. 548.—Eliason, 1920, p. 63.—Fauvel, 1927, p. 154, fig. 55, a-h.— 
Augener, 1928, p. 749.—Monro, 1930, p. 163.—Annenkova, 1934, p. 322; 
1937, p. 179; 1938, p. 189.—Hartman, 1942a, p. 69; 1944a, pp. 334, 341, 
pl. 34, fig. 8; 1945, p. 37; 1947, p. 404, pl. 43, figs. 1, 2; 1948, p. 41; 1951, 
p. 101.—Berkeley and Berkeley, 1948, p. 129; 1952, p. 100, figs. 206-208.— 
Gorbunov, 1946, p. 39.—Thorson, 1946, p. 110, fig. 60.—Zatsepin, 1948, 
p. 138, pl. 33, fig. 14—Hartman and Reish, 1950, p. 40.—Wesenberg-Lund, 
1950a, p. 39; 1950b, p. 91; 1951, p. 84. 

Capitella capitata var. antarctica Monro, 1930, p. 164. 


Description.—Length 23 mm., width 1 mm. (up to 120 mm.— 
Augener, 1928). Body attenuated slightly anteriorly, much more 


so posteriorly, fragile. Prostomium and achaetous buccal segment 
fused, forming wide obtuse cone, somewhat flattened dorsoventrally, 


MARINE POLYCHAETE WORMS—PETTIBONE 299 


usually concave dorsally," with"pair of small ventral eyes and pair of 
crescent-shaped nuchal slits behind eyes (best seen from lateral view). 
Thoracic region smooth, without reticulated integument, consisting 
of first 9 setigerous segments, each of first 7 with four bundles of 
capillary setae which on setiger 7 may be only capillaries, only crotch- 
ets, or both (in young specimens, crotchets may be present from 
setiger 4). Setigers 8 and 9 modified, having neuropodia with row of 
hooded crotchets, and with large, recurved, amber-colored acicular 
genital hooks located middorsally between these two segments. 
Abdominal segments with slightly raised dorsal and ventral para- 
podial tori each bearing a row of hooded crotchets. Without spe- 
cialized branchiae or lateral organs. Pygidium a short ring, may be 
inflated. Conor: In alcohol: Without color or reddish brown. 
Females with large yolky eggs (Point Barrow, August 9, 30, 1949; 
September 1, 6, 1949.) 

Remarks.—According to Fauvel and others, in C. capitata the genital 
hooks on setigers 8 and 9 are present only in the males, the females 
having only crotchets. On all the specimens from Point Barrow, 
genital hooks are present, including some females containing large 
yolky eggs. They agree in all other respects with C. capitata. Having 
genital hooks in both males and females, they would fall under the 
very closely related Capitellides Mesnil. They differ from C. giardi 
Mesnil in the larger number of genital hooks. They are herein re- 
ferred to the cosmopolitan C. capitata with the tentative idea that the 
relative development of genital hooks in the female may prove to be 
a variable character. 

New records—Arctic AuaskKa: Eluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 7.5 miles from 
shore, 23.3-70 fms., on bottoms of stones and various combinations 
of gravel, stones, rocks, large perforated rocks (8 stations, 26 speci- 
mens). Brrina Sxa: St. Paul Island, Pribilofs, in holdfasts with 
Fabricia sabella and lumbrinerids, W. L. Hahn, 1911. West Coast 
Norra America: Corona del Mar, California, from branchial cavity 
of crab, Loxorhynchus grandis, MacGinitie, 1948. East Coast Nortu 
America: Off Labrador, 6 fms., mud-sand, Blue Dolphin Expedition, 
1949; off Massachusetts, 15 fms., U. S. Fish Commission; Woods Hole 
region, Massachusetts, Pettibone, 1951. 

Distribution.— Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Novaya 
Zemlya, KaraSea. Also Iceland, Faroes, Norway to France, Madeira, 
Mediterranean, Black Sea; Hudson Bay to North Carolina, Texas; 
Bering Sea to southern California; north Japan Sea. Southern lati- 
tudes: Magellan Straits, South Georgia, Bouvet Island, South Africa, 
Kerguelen. In low water to 500 fathoms. 

261112547 


300 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Family ARENICOLIDAE 


Body elongate, cylindrical, integument reticulated and divided into 
secondary annuli. Body divided into two or three distinct regions: 
(1) anterior, abranchial; (2) abdominal, branchial, each segment with 
pair of dorsally-borne branched gills; (8) caudal, achaetous and 
abranchial; this region may be lacking. Prostomium small, simple 
or trilobed, without appendages; with nuchal groove (fig. 34, a). 
Parapodia biramous; notopodia conical, setigerous, with spiny, capil- 
lary setae; neuropodia with sigmoid crotchets in semicircular tori. 
Without dorsal, ventral, and anal cirri. Anus terminal. Proboscis 
eversible, globular, papillate, unarmed. Constructs tubes or burrows. 

Represented by a single species from Point Barrow. 


Genus Arenicola Lamark, 1801 
Arenicola glacialis Murdoch, 1884 


FIGURE 34, a 


Arenicola glacialis Murdoch, 1884, p. 522; 1885, p. 155.—Ashworth, 1910, p. 24, 
figs. 10-14; 1912, p. 111, figs. 46, 47, pl. 6; 1924, p. 3, fig. 1. 

Description.—Length up to 205 mm., width 11 mm., 17 setigerous 
segments. Body cylindrical, tapering slightly anteriorly and gradu- 
ally posteriorly. Prostomium small, may be pretty well retracted in 
nuchal groove; median lobe small, lateral lobes larger, curved (fig. 
34, a). Body divided into three regions: (1) anterior region of six 
abranchial setigers, notopodia conical with spiny capillary setae, 
neuropodia low tori with few sigmoid crotchets; (2) branchial region 
of 11 setigers each divided into five superficial annuli, with short, 
bushy, arborescent branchiae at base of conical notopodia, with gill 
axes short, branching one or two times or simple; neuropodia raised 
tori with long row of crotchets; on more posterior segments, tori 
approach midventral line; (3) caudal achaetous and abranchial region 
with papillae feebly developed, without processes (as in A. cristata). 
Nephridial pores six pairs, on setigers 4-9, posterior to upper part of 
neuropodial tori. Pair of statocysts on peristomium, open to exterior, 


Ficure 34.—Arenicolidae: a, Arenicola glacialis, prostomium. Maldanidae: b, Praxillella 
praetermissa, ventral view anterior end; c, same, frontal view cephalic plate; d, same, 
pygidial funnel; ¢, same, posterior end; f, same, crotchet from neuropodium; g, Maldane 
sarsi, lateral view anterior end; h, same, lateral view posterior end; i, Nicomache lum- 
bricalis, lateral view anterior end; 7, same, lateral view posterior end; k, Nicomache 
personata, lateral view posterior end; J, Petaloproctus tenuis, lateral view anterior 
end; m, same, lateral view posterior end. Sabellariidae: n, Idanthyrsus armatus, dorsal 
view anterior end; 0, same, lateral view anterior end; p, same, palea from inner row; 
g, same, palea from outer row. (For explanation of symbols, see p. 210.) 


MARINE POLYCHAETE WORMS—PETTIBONE 301 


P 


FicuRE 34.—For explanation see facing page. 


302 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


with numerous sand grains as statoliths. Cotor: In life: Deep orange 
to brownish red. In alcohol: Bluish to grayish black or brown. 

New records.—Anrctic ALASKA: Point Barrow base, washed ashore 
August 21, 24, 28, September 6, 12, 19, 22, October 16, 1949 (54 
specimens) ; off Point Barrow base, along shore, 1.7 fms., on bottom 
of gravel with mud (2 stations, 3 specimens). 

Distribution.—Alaskan and Canadian Arctic. In 1.7—-3 fathoms. 


Family MALDANIDAE 


Body cylindrical, truncated at both ends, without differentiated 
regions, segments usually few, some much longer than wide. Pro- 
stomium a small cephalic crest fused with achaetous buccal segment, 
forming ovoid head, without appendages, with or without oblique 
limbate brim or plate (fig. 34, 6, 7, 2); pair of nuchal slits on either side 
cephalic crest. In anterior region with numerous glandular cells, 
diffused or grouped in clear bands alternating with pigmented rings. 
Parapodia biramous; notopodia with capillary setae; neuropodia with 
projecting tori bearing rows of crotchets (fig. 34, f) with dentate ros- 
trum and rostral hairs; the crotchets may be replaced by few stout 
acicular spines on first few setigers. Without dorsal or ventral cirri; 
usually without branchiae. Pygidium a limbate plate, spatulate or 
funnellike (fig. 34, h, 7, m). Proboscis eversible, unarmed, globular. 
Live in tubes in sand or mud; tube cylindrical, membranous, covered 
with thin to thick layer of mud, sand, gravel, more or less resistant. 

Represented by four genera and five species from Point Barrow. All 
the genera are without specialized collarettes, without special glandular 
ventral shield on setiger 8, with neuropodial crotchets in single rows. 


Key to the genera of Maldanidae from Point Barrow 


1. ‘Head. with limbate plate: (fig. 34, b, g)...-L---=22£2..__. 22 eee 2 
Head without limbate plate (fig. 34, 7, J). With 1-2 stout acicular setae in 
neuropodia of first 3 setigers. Pygidium funnel-shaped or forming concave 
plate, with or without cirri; anus opening on surface of plate__________- 3 
2. Head with low cephalic crest, bordered by oblique limbate plate, entire or 
notched laterally (fig. 34, b,c). With ventral crotchets beginning on setiger 1. 
Pygidium funnel-shaped, bordered with cirri, with anus in center of somewhat 
protruding anal cone within the funnel (fig. 34, d)______ Praxillella (p. 303) 
Head with middorsal convex cephalic crest or keel, bordered by limbate plate, 
divided into 3 parts by 2 deep lateral grooves (fig. 34, g). Without neuro- 
setae on setiger 1; crotchets begin on setiger 2. Pygidium a slightly oblique 
oval plate, bordered by rim which is incised laterally, smooth or denticled; 
anus dorsal to pygidial dise (fig. 34, fi)... 22 - 2-22-22 Maldane (p. 303) 

3. Pygidium slightly oblique, funnel-shaped, bordered with cirri (fig. 34, 7, k). 
Nicomache (p. 304) 
Pygidium an asymmetrical, foliaceous, concave plate, reduced dorsally, wider 
ventrally, with border smooth or crenulate (fig. 34, m) -Petaloproctus (p. 306) 


MARINE POLYCHAETE WORMS—PETTIBONE 303 


Genus Praxillella Verrill, 1881 
Praxillella praetermissa (Malmgren, 1865) 
Figure 34, b-f 


Prazilla praetermissa Malmgren, 1865, p. 191; 1867, p. 100, pl. 11, fig. 62. 

Prazillella praetermissa Verrill, 1881, pp. 298, 305, 309, 312—Webster and 
Benedict, 1887, p. 746.—Arwidsson, 1907, p. 192, figs—Eliason, 1920, p. 
66.—Augener, 1928, p. 762.—Annenkova, 1937, p. 182; 1938, p. 194.— 
Zatsepin, 1948, p. 142, pl. 35, fig. 1—Wesenberg-Lund, 1948, p. 41, figs. 
20, 21; 1950a, p. 43; 1950b, p. 98; 1951, p. 93. 

Clymene (Prazillella) praetermissa Fauvel, 1927, p. 179, fig. 62, a-e. 


Description —Length 45 mm., width 1 mm. (up to 120 mm. long, 
2.5mm. wide—Malmeren, 1865), 19 setigerous segments, 4-5 achaetous 
segments with rudimentary parapodia. Head with oblique limbate 
plate, with border entire, slightly or deeply incised laterally, slightly 
incised posteriorly, terminating anteroventrally in obtuse cone (not 
produced into digitiform process) ; median cephalic crest long; nuchal 
grooves one-half to two-thirds its length. Neuropodia of anterior 
three setigers with 2-10 slightly modified crotchets. Pygidial funnel 
with flat floor, with projecting anal cone with larger valvule on ventral 
side of cone; border of funnel with 20-38 subequal papillae except 
for a longer ventral one. Proboscis papillate basally. Conor: In 
life: Yellow with wide, red and white bands on some anterior segments. 
In alcohol: Brownish to gray, with whitish bands anterior to setae 
on setigers 4-8. Tuss: Of clay, sand, small bits of rock; may be quite 
resistant. 

New records—Arctic AtasKa: Off Point Barrow base, up to 7.5 
miles from shore, 20-36 fms., on various combinations of mud, sand, 
gravel, stones, rocks, large perforated rocks, shells (7 stations, 10 
specimens). East Coast Norru America: Off Labrador, 5-8 fms., 
Blue Dolphin Expedition, 1949; off Cape Cod, Massachusetts, 14-19 
fms., U. S. Fish Commission. 

Distribution —Widely distributed in the Arctic: Alaskan Arctic, 
Davis Strait, Greenland, Spitsbergen, White Sea, Novaya Zemlya, 
Kara Sea. Also Iceland, Faroes to English Channel, Mediterranean; 
Labrador to Massachusetts; north Japan Sea. In 7-1,111 fathoms. 


Genus Maldane Grube, 1860 
Maldane sarsi Malmgren, 1865 
FieureE 34, g, h 


Maldane sarsi Malmgren, 1865, p. 188; 1867, p. 99, pl. 10, fig. 57.—Arwidsson, 
1907, p. 251, figs —Eliason, 1920, p. 66.—Fauvel, 1927, p. 197, fig. 69, a-i; 
1932, p. 202.—Augener, 1928, p. 759.—Annenkova, 1937, p. 179; 1938, p. 
192.—Okuda, 1939, p. 239.—Berkeley and Berkeley, 1942, p. 199.—Hartman, 


304 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


1944a, pp. 335, 342, pl. 33, figs. 3, 4; 1951, p. 106.—Zatsepin, 1948, p. 142, 
pl. 33, fig. 3.— Wesenberg-Lund, 1948, p. 48, figs. 24, 25; 1950a, p. 44; 1950b, 
p. 100; 1951, p. 94.—Hartman and Reish, 1950, p. 37. 

Maldane sarst var. antarctica Arwidsson, 191la, p. 32, figs—Munro, 1930, p. 
169; 1936, p. 168. 

Maldane sarsi var. Moore, 1923, p. 237. 

Maldane sarsi var. tropica Monro, 1937, p. 307, fig. 24. 

Description.—Length up to 110 mm., width 3.9 mm., setigerous 
segments 19, two antanal achaetous segments. Head with long, 
convex keel, ending in blunt process above mouth, bordered by limbate 
plate, with border smooth, usually incised laterally, rather high pos- 
teriorly; nuchal organs short, divergent; without ocelli. Anterior 
segments biannulate. With notosetae only on first setiger; neuro- 
podial crotchets begin on setiger 2. Two antanal segments short, 
with achaetous tori slightly marked. Pygidium an oblique flat, oval 
plate, with border usually slightly incised laterally, smooth or slightly 
crenulate ventrally. Anus a folded cushion dorsal to and close to 
pygidial rim. Tuse: Free, circular, thin to thick walled, of fine 
agglomerated mud and sand grains. 

New Records—Arctic ALtAsKA: Off Pomt Barrow base, 3 miles 
from shore, 20.3 fms., on bottom of mud, gravel, stones, shells (1 
station, 1 specimen). Burrine Sa: Albatross Sta. 3603, 55°23’ N., 
170°31’ W., 1,771 fms., 1895. Canapian Arctic: Northwest shore 
Daniels Island, mouth Newell Sound, Frobisher Bay, Baffin Land, 
10-30 fms., Bartlett. East Coast Norra America: Off Labrador, 
30 fms., Blue Dolphin Expedition, 1950; off Nova Scotia, Maine, 
Massachusetts, Rhode Island, 14-487 fms., U. S. Fish Commission., 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan 
and Canadian Arctic, Davis Strait, Greenland, Spitsbergen, Novaya 
Zemlya, Kara Sea. Also Iceland, Norway to France; Labrador to 
Rhode Island, Louisiana, Mississippi, eastern Texas; Bering Sea to 
southern California; north Japan Sea to Japan; South Arabian coast 
(var. tropica); Bay of Bengal. Southern latitudes: South Georgia, 
South Shetlands, Palmer Archipelago (var. antarctica) ; south Australia. 
In low water to 1,771 fathoms. 


Genus Nicomache Malmgren, 1865 


Both species have buccal segment fused with prostomium forming 
hood-shaped head, irregularly rounded, with thickened anteroventral 
border, forming projecting upper lip (fig. 34,7). First three setigerous 
segments each with one or two stout, acicular, spinelike neurosetae. 
Some of notopodia with long, hairlike sinuous dorsal setae. Pygidium 
with anus terminal in center of short, slightly oblique funnel, with 
papillae on border of funnel (fig. 34, 7, k). 


MARINE POLYCHAETE WORMS—PETTIBONE 305 


Key to the species of Nicomache from Point Barrow 


1. Single achaetous antanal segment (fig. 34, k)_______---------- N. personata 
Two achaetous antanal segments (fig. 34, j).-.._____-------- N. lumbricalis 


Nicomache lumbricalis (Fabricius, 1780) 
Ficure# 34, 1,7 


Sabella lumbricalis Fabricius, 1780, p. 374. 

Nicomache lumbricalis Malmgren, 1867, p. 99, pl. 10, fig. 60.—Webster and 
Benedict, 1884, p. 731; 1887, p. 745.—Arwidsson, 1907, p. 86, pl. 8, figs. 
244, 245.—Fauvel, 1927, p. 190, fig. 66, a-i—Augener, 1928, p. 764.—Monro, 
1930, p. 173.—Annenkova, 1934, p. 322; 1937, p. 181; 1938, p. 193.— Hartman, 
1944a, pp. 335, 342; 1948, p. 42.—Zatsepin, 1948, p. 144, pl. 26, fig. 1.— 
Wesenberg-Lund, 1948, p. 23, figs. 10, 11; 1950a, p. 41; 1950b, p. 95; 1951, 
p. 90.—Berkeley and Berkeley, 1952, p. 54, figs. 111, 112. 

Nicomache carinata Moore, 1906b, p. 242, pl. 11, figs. 36-39, pl. 12, figs. 48, 44; 
1923, p. 227. 

Nicomache lumbricalis var. borealis Arwidsson, 1907, p. 94, figs.; 1922, p. 6.— 
Eliason, 1920, p. 65. 


Description —Length 40-60 mm., width 2.5-3 mm., setigerous 
segments 21-22, two short achaetous antanal segments (Point Barrow 
and Labrador specimens with 22 setigers; type of N. carinata with 21; 
up to 160 mm. long, 5 mm. wide—Fauvel, 1927). Prostomium with- 
out ocelli; nuchal organs S-shaped. Pygidial funnel with 14-30 
subequal, pointed, triangular papillae. Conor: In alcohol: Reddish 
brown on sides and dorsal surface of head and first three setigers. 
TusBeE: Sandy, with small pieces of rock, with smooth membranous 
lining (Point Barrow), or thick, sandy, and coiled (Labrador; small 
ones on terebellid tubes). 

Remarks.—They may share their tubes with the commensal poly- 
noid Hnipo gracilis Verrill (=E. cirrata Treadwell) reported by Berke- 
ley and Berkeley from Alaska; also from off Nova Scotia and Mas- 
sachusetts (in USNM). 

New records.—Arctic Auaska: Off Point Barrow, up to 12.1 miles 
from shore, 24.7-123.5 fms., on bottoms of mud, worm tubes, stones, 
and various combinations of rocks, stones, gravel (6 stations, 18 
specimens). Brrina Sea: Cape Prince of Wales, 23 fms., Dall, 1874. 
Albatross Sta. 3311, 53° 59’ N., 166° 29’ W., 85 fms., and Sta. 3313, 
54° 01’ N., 166° 27’ W., 68 fms., 1890. AxasKa: Albatross Sta. 2848, 
Unalaska to Cook Island, 1888; Eastern Harbor, Sitka, 15 fms., Dall, 
1874. KamcnatKa: Albatross Sta. 3644, 51° 09’ N., 157° 48’ W., 96 
fms., 1896. East Coast Norta America: Off Labrador, 30-125 
fms., Blue Dolphin Expeditions, 1949, 1950, 1951; off Nova Scotia, 
Bay of Fundy, Maine, New Hampshire, Massachusetts, 26-150 fms., 
U.S. Fish Commission. 

Distribution—Widely distributed in the Arctic: Siberian and 


306 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


Alaskan Arctic, Greenland, Jan Mayen, Spitsbergen, White Sea, 
Novaya Zemlya, Kara Sea. Also Bering Sea to Lower California; 
north Japan Sea; Iceland, Norway to English Channel, Santander; 
Labrador to Massachusetts; South Africa. In low water to 1,400 


fathoms. 
Nicomache personata Johnson, 1901 


Figure 34, k 


Nicomache personata Johnson, 1901, p. 419, pl. 13, figs. 134-139.—Arwidsson, 
1922, p. 7, pl. 1, figs. 5, 6—Hartman, 1948, p. 41, fig. 11, d~-g.—Berkeley 
and Berkeley, 1952, p. 54, figs. 109, 110. 

Nicomache minor Arwidsson, 1907, p. 100, pl. 2, figs. 68-73; pl. 8, figs. 252-256.— 
Augener, 1928, p. 765.—Annenkova, 1937, p. 181; 1938, p. 193. 

Nicomache maculata Arwidsson, 1911b, p. 209, pl. 18, figs. 13-19; pl. 19, figs. 
27-30.—Fauvel, 1927, p. 191, fig. 66, k-r. 

Description.—Up to 110 mm. long, 2 mm. wide, 22 or 23 setigerous 
segments, single achaetous antanal segment (Point Barrow specimen 
with 23 setigers, Puget Sound specimen with 22). Prostomium with 
numerous ocelli (may be absent) ; nuchal clefts short, arched. Pygidial 
funnel with 16-26 papillae which are short, subequal, triangular, or 
very unequal, some being longer, some very short, few may be bifid 
or trifid. Co tor: In alcohol: Without color, or brownish, especially 
onanteriorend. Tuss: Thick walls of coarse sand particles cemented 
together. 

New records—Arctic Ataska: Off Point Barrow base, up to 8 
miles from shore, 21.7—75.5 fms., on various combinations of pebbles, 
rocks, gravel, stones (3 stations, 5 specimens). Brrine Sra: Alba- 
tross Sta. 3289, 56° 44’ N., 159° 16’ W., 16 fms., 1890. Wersr Coast 
Norta America: Security Bay, Alaska, Jones; Washington and Puget 
Sounds, low water to 61 fms., Pettibone. 

Distribution.—Arctic Alaska, Spitsbergen; Bering Sea to Washing- 
ton; north Japan Sea; Scandinavia, Ireland, North Sea to English 
Channel. In low water to 75.5 fathoms. 


Genus Petaloproctus Quairefages, 1865 


Petaloproctus tenuis (Théel, 1879) 


Figure 34, 1, m 


Maldane tenuis Théel, 1879, p. 57, pl. 4, figs. 52-54. 

Maldane filifera Verrill, 1879, p. 179. 

Petaloproctus tenuis Arwidsson, 1907, p. 114, pl. 6, fig. 190a.—Annenkova, 1937, 
p. 181; 1938, p. 193.—Zatsepin, 1948, p. 145, pl. 34, fig. 5-—Wesenberg-Lund, 
1948, p. 31.—Berkeley and Berkeley, 1952, p. 55, fig. 113. 

Petaloproctus tenuis var. borealis Arwidsson, 1907, p. 118, pl. 3, figs. 85-90; pl. 8, 
figs. 268-272.—Eliason, 1920, p. 66.—Hartman, 1948, p. 42.—Berkeley and 
Berkeley, 1952, p. 56, figs. 114-116. 

Petaloproctus filifer Hartman, 1942b, p. 131, fig. 11, a-b. 


MARINE POLYCHAETE WORMS—PETTIBONE 307 


Description —Length 18 mm., width 0.5 mm., 20-21 setigerous 
segments, single achaetous antanal segment (may appear as two or 
three on longer ventral side). Prostomium rounded, with convex 
median ridge and anteroventrally extended brim. First three setigers 
with one or two acicular neurosetae; with some long, threadlike, sinu- 
ous notosetae covered with spinules. Pygidium with oblique, slightly 
concave, disclike, asymmetrical plate (absent dorsally), wide on ven- 
tral side, with rim smooth or slightly crenulate; anal opening terminal. 

Remarks.—The Point Barrow specimen agrees with the stem form 
of P. tenuis, with 20 setigerous segments and the rim of the pygidial 
disc slightly crenulate (in var. borealis and filifer there are 21 setigers 
and the rim of the pygidial disc is smooth). 

New records —Arctic ALasKa: Off Point Barrow base, 16 miles 
from shore, 78.2 fms., on bottom of worm tubes, few rocks (1 station, 
1 specimen). Canapran Arctic: Ducketts Cove, Hurd Channel, 
Melville Peninsula, 1-14 fms., in sandy tubes on empty tubes of 
Pectinaria, Bartlett, 1933. GREENLAND: Small bay, Camp No. 2, 
Nugsuak Peninsula, 74°7’ N., on empty tubes of Pectinaria, J. C. 
Martin, 1897. 

Distribution. Scattered records in the Arctic: Alaskan and Cana- 
dian Arctic, East Greenland, Spitsbergen, Novaya Zemlya. Also off 
Cape Cod, Massachusetts; southeastern Alaska to British Columbia; 
north Japan Sea. In 1-833 fathoms. 


Family SABELLARIIDAE 


Body divided into three regions: (1) anterior thoracic region includ- 
ing two anterior thoracic segments with parapodia reduced to neuro- 
podia with capillary setae and two large, more or less fused, heavy 
columns, the opercular peduncles, directed anteriorly, bearing very 
modified setae arranged in one to three concentric rows and forming an 
operculum for closing opening of tube; three or four parathoracic 
biramous segments with capillary neurosetae and paddlelike or 
styliform notosetae and dorsal falciform branchiae (fig. 34, n—q); (2) 
abdominal region with notopodia in form of broadly flattened pinnules 
bearing pectinate uncini, and neuropodia with capillary setae; with 
dorsal, simple, ligulate or falciform branchiae; (3) caudal region, 
achaetous, apodous, cylindrical, without appearance of segmentation. 
Prostomium indistinct, between opercular peduncles, with pair of 
palps and usually a median tentacle. Mouth unarmed, usually sur- 
rounded with numerous filiform tentacles. Tube of sand or fine 
gravel, concreted, resistant, may be solitary or colonial, constructing 
sandy reefs. 

Represented by a single species from Point Barrow. 


308 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Genus Idanthyrsus Kinberg, 1867 
Idanthyrsus armatus Kinberg, 1867 


Figure 34, n-q 


Idanthyrsus armatus Kinberg, 1867, p. 350.—Johansson, 1926, p. 9; 1927, p. 90.— 
Monro, 1930, p. 117, fig. 73; 1936, p. 172.—Annenkova, 1937, p. 184; 1938, 
p. 196.—Okuda, 1938a, p. 242, figs. 4, 5—Hartman, 1944c, p. 336, pl. 31, 
fig. 36.—Berkeley and Berkeley, 1952, p. 107, figs. 220-222. 

Idanthyrsus ornamentatus Chamberlin, 1919, p. 262, pl. 3, figs. 2-5—Hartman, 
1944¢, p. 337, pl. 31, fig. 34; 1948, p. 43.—Hartman and Reish, 1950, p. 41. 

Description.—Length up to 60 mm., width 5 mm., caudal lobe 11 
mm. Body widest anteriorly, tapering gradually posteriorly, with 
achaetous caudal region sharply set off from body. Opercular pe- 
duncles elongated, semicylindrical, completely separated and diver- 
gent anteriorly, obliquely truncated distally, with two rows modified 
golden setae or paleae; inner row paleae rather stout, smooth, gently 
curved, tapering gradually to acute tips, 7-16 per column; outer row 
paleae more slender, straight or slightly curved, with coarse lateral 
spines, 16-36 per column; row of 9-15 distal papillae per column, at 
base of outer paleae. One or two pairs heavy, deep amber-colored, 
strongly-bent nuchal hooks on dorsal side opercular peduncles. 
Numerous filiform oral tentacles on ventral side opercular peduncles. 
Pair of grooved slender palps and median tentacle between bases of 
peduncles. First thoracic segment short, with ventral group capillary 
setae lateral to conical papilla. Second segment with lateral group 
capillary setae, three papillae and dorsal branchiae. Three para- 
thoracic segments with rectangular palletlike pinnules with flattened, 
paddlelike notosetae; neurosetae bipinnate capillary setae. Abdom- 
inal segments with wide, flattened dorsal pinnules bearing pectinate 
uncini; capillary barbed neurosetae. Caudal lobe smooth, achaetous. 
Paired dorsal branchiae cirriform, with transverse ridges on medial 
side, begin on setiger 2, absent on last few abdominal segments. 
Cotor: In life: Reddish brown to dark violet anteriorly, branchiae 
red, paleae golden yellow. In alcohol: Colorless, with purple 
patches anteriorly. Tus: Thick, of coarse sand grains neatly ce- 
mented together, very resistant, one side flattened, attached to rocks, 
shells, crustaceans; upper surface convex. 

Remarks.—I. ornamentatus Chamberlin is herein referred to J. 
armatus, following Okuda (1938). According to observations by G. 
E. MacGinitie it is a hardy species, a specimen living in a pail in the 
laboratory for three days, on a rock covered with a mass of barnacles, 
etc. Male specimen, collected from 70 fathoms, spawned in the 
laboratory August 9, 1949. 


MARINE POLYCHAETE WORMS—PETTIBONE 309 


New records—Arctic AuasKa: Off Point Barrow base, up to 7 
miles from shore, 4.3—70 fms., on bottoms of mud, and gravel, stones 
(3 stations, 6specimens). Brrine Sna: Bering Island, N. Grebintzky, 
1884. Wurst Coast Norts America: Albatross Sta. 2842, 54°15’ N.., 
166°3’ W., 72 fms.; Washington and Puget Sounds, 10-110 fms., 
Pettibone. 

Distribution —Arctic Alaska, Bering Sea to Panamé; north Japan 
Sea to Japan; west coast South America (Chile), Straits of Magellan, 
Falkland Islands. In low water to 110 fathoms. 


Family STERNASPIDAE 


Body very short, grublike, aberrant; segments few, uniramous or 
achaetous, without parapodial lobes (fig. 35, a). Prostomium re- 
duced to small rounded tubercle, without appendages. Mouth 
rounded, subterminal, ventral to prostomium. Body divided into 
three regions: (1) anterior, with complete rings, with lateral concentric 
rows of strong acicular yellow setae on anterior three segments 
(anterior region may be retracted into more posterior segments); 
(2) middle, with segmental divisions absent on midventral part, 
without setae visible externally; (3) posterior, with ventral horny 
shield formed by two trapezoidal plates provided on their external 
sides with radiating bundles of stiff, barbed and smooth capillary 
setae (fig. 35, 6). Numerous, long, filiform anal branchiae, inserted 
on posterior end. Anus terminal. Lives in mud. 

Represented by a single species from Point Barrow (the family is 
usually considered to have a single cosmopolitan species). 


Genus Sternaspis Otto, 1821 
Sternaspis scutata (Ranzani, 1817) 
Figure 35, a, b 


Thalassema scutata Ranzani, 1817, p. 1457, pl. 11, figs. 10-13. 

Sternaspis fossor Stimpson, 1854, p. 29, pl. 2, fig. 19——Webster and Benedict, 
1884, p. 725.—Moore, 1909b, p. 144; 1923, p. 218.—Hartman, 1944a, pp. 336, 
342, pl. 33, fig. 15.—Berkeley and Berkeley, 1952, p. 59, fig. 123. 

Sternaspis islandica Malmgren, 1867, p. 87, pl. 14, fig. 85. 

Sternaspis scutata Moore, 1903, p. 487; 1908, p. 357; 1923, p. 218.—Fauvel, 1927, 
p. 216, fig. 76, a-g; 1932, p. 213; 1933, p. 52; 1934a, p. 60.—Monro, 1930, 
p. 178.—Annenkova, 1937, p. 185; 1938, p. 196.—Hartman, 1942c, p. 102.— 
Gorbunov, 1946, p. 39.—Wesenberg-Lund, 1949, p. 345; 1950a, p. 46; 1950b, 
p. 104; 1951, p. 98.— Hartman and Reish, 1950, p. 38. 

Sternaspis scutata var. africana Monro, 1930, p. 179. 


310 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Description.—Up to 31 mm. long, 14 mm. wide. Body variable in 
shape; may be inflated at both ends, anterior region may be retracted 
into more posterior segments. Integument densely pilose, covered 
with fine filiform papillae. Anterior region of seven segments; 
segments 1 and 5-7 achaetous; segments 2—4 each with lateral semi- 
circular row of strong, short, yellow setae which are arched, pointed 
to blunt (worn down), diminishing in size toward ventral side; pair 
of long genital papillae on segment 7. Middle region of six segments, 
well defined except for narrow midventral strip, with bundles of capil- 
lary setae embedded in body wall. Posterior region with five or more 
segments, with ventral horny trapezoidal plates showing concentric 
and radiating striae, each plate with 16 radiating bundles of stiff 
setae, 9 on lateral and 7 on posterior borders. Anal filaments or 
branchiae very numerous, long, filiform, sometimes spiralled. Conor: 
In alcohol: Without color or brownish, with rusty-red ventral shield. 

New records.—Arctic AutasKa: Off Point Barrow base, up to 5 
miles from shore, 1.7-24.7 fms., on bottoms of mud and gravel with 
mud (3 stations, 5 specimens). Brrine SEA: 63°37’ N., 165°19’ W., 
12 fms., Stoney, 1884; off mouth Yukon River, 3 fms., E. W. Nelson, 
1877. AwasKa: Iluiliuk Harbor, Unalaska, Dall, 1871; Albatross 
Sta. 3340, 55°26’ N., 155°26’ W., 695 fms., and Sta. 3330, 54° N., 
166°53’ W., 351 fms., 1890. Chiachi Islands, 20 fms., 1874; off 
Round Island, Coal Harbor, Unga Island, 6-8 fms., 1872; Eastern 
Harbor, Sitka, 15 fms., gravelly-mud, 1874; all collected by Dall. 
West Coast Norta America: Washington and Puget Sounds, 
10-111 fms., mud, Pettibone. East Coast Nortn America: Nova 
Scotia, Maine, Massachusetts, 5-102 fms., U. S. Fish Commission. 

Distribution.—Scattered records in the Arctic: Siberian and Alaskan 
Arctic, Greenland. Also Iceland to English Channel, Mediterranean, 
Adriatic, Gulf of Oman, Persian Gulf; Nova Scotia to Massachusetts; 
Bering Sea to Panamé4; north Japan Sea to Japan, China. Southern 
latitudes: South Shetlands, Palmer Archipelago, South Orkneys, 
South Georgia, west Africa, Indian Ocean, Australia, New Zealand. 
In 1.7-766 fathoms. 


Family PEcTINARIIDAE (AMPHICTENIDAE) 


Body short, conical, with segments few in number. Prostomium 
indistinct, fused with buccal segment forming a truncate anterior end 
provided with two bundles of paleae (large, flattened, golden setae) 
in a horizontal row directed obliquely anteriorly, forming an operculum 
for the tube or as a fork for digging in sand or mud; ventral to paleae, 
an antennular or tentacular membrane bordered or not with short to 
long filiform papillae, surrounding numerous prehensile oral tentacles 


MARINE POLYCHAETE WORMS—PETTIBONE 311 


not retractile in mouth; dorsal and posterior to paleae a semilunar 
flat or concave cephalic plate limited posteriorly by an entire or 
crenulate rim; two pairs tentacular cirri with first pair lateral to base 
of paleae, second pair lateral to rim of cephalic plate (fig. 35, ¢, d). 
Body divided into three regions: (1) anterior thoracic, including two 
achaetous segments each with a pair of lateral pectinate branchiae, 
three notopodial uniramous setigers lacking neuropodia; ventral side 
anterior region with thick glandular cushions; (2) median abdominal, 
with biramous segments (except last few may have notosetae only), 
notopodia with setae capillary, limbate, smooth or finely denticled; 
neuropodia in form of wide pinnules bearing single row of pectiniform 
uncini; (3) posterior anal plaque or scapha, small, foliaceous, concave 
dorsally, folded under abdomen, with rudimentary achaetous segments 
except for the first segment which has a series of acicular, more or 
less recurved setae (the scaphal hooks); an oval or filiform ligule 
above the anus (fig. 35, g, h). Tube free, conical, rigid, fragile, open 
at both ends, straight or arcuate, formed of a single layer of cemented 
sand grains or shells, and lined by a membrane. Live in sand or mud, 
with anterior and larger end of tube directed below. May secrete 
mucus in abundance. 

Represented by a single genus (and subgenus) and two species 
from Point Barrow. 


Genus Pectinaria Lamark, 1818 
Subgenus Cistenides Malmgren, 1865 


Both species have the tube slightly arched, lined with rusty- 
colored membrane. Tentacular membrane free, the lateral portions 
not fused to paleal segment, with 30-45 filiform to clavate marginal 
papillae. Oral tentacles short, thick, grooved. Dorsal rim of flat 
paleal segment entire, not denticled (fig. 35, d). Usually 17 segments 
with capillary notosetae (15-17, the last two segments may have the 
notosetae few in number or lacking), of which 12 (beginning on setiger 
4) have elongate flattened pinnules bearing single rows of neuropodial 
uncini; pectiniform uncini thin, flat, with 3-4 major teeth and 4-5 
small ones above basal process (fig. 35, f). Anal plaque distinctly 
separated from abdomen by constriction, suboval, festooned laterally 
(rudimentary parapodia) ; basally with 7-11 pairs short, heavy scaphal 
hooks (fig. 35, g). 


Key to the species of Pectinaria (Cistenides) from Point Barrow 


1. Cephalic paleae usually 7-9 pairs (7-13), heavy, with blunt tips (or very short 
eee GLA Get fe Ty ek ee ee ok ee P. (C.) granulata 
Cephalic paleae usually 11-14 pairs (9-15), long, tapered to slender pointed 
GEES EL ON Sra A ee ee ne ne kne Seeeeeinn So LK ghd P. (C.) hyperborea 


312 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 
Pectinaria (Cistenides) granulata (Linné, 1767) 
FiaureE 35, i-k 


Sabella granulata Linné, 1767, p. 1268. 

Cistenides granulata Malmgren, 1865, p. 359.—Webster and Benedict, 1887, p. 
747.—Chamberlin, 1920, p. 25.—Berkeley and Berkeley, 1942, p. 200.— 
Hartman, 1944a, pp. 335, 342.— Wesenberg-Lund, 1950b, p. 105; 1951, p. 99. 

Pectinaria brevicoma Johnson, 1901, p. 423, pl. 15, figs. 151-156.—Not Moore, 
1923, p. 216 (=P. californiensis Hartman; in USNM). 

Pectinaria (Cistenides) granulata Hessle, 1917, p. 77.—Nilsson, 1928, p. 28, fig. 
8.—Annenkova, 1937, p. 186; 1938, p. 198.—Treadwell, 1937, p. 32. 

Cistenides brevicoma Hartman, 1941b, p. 331, pl. 50, figs. 13, 14, 16; pl. 52, fig. 
23; 1944b, p. 268. 

Pectinaria (Cistenides) brevicoma Berkeley and Berkeley, 1952, p. 106, figs. 218, 
219. 


Description — Body 20-52 mm. long, 6-8 mm. wide. Tube 38-52 
mm. long, 7-8 mm. in greatest diameter, of rather coarse sand grains, 
nearly uniform in size. Paleae usually 8 or 9 pairs (7-13), short, 
heavy, with tips blunt (may have short, sharp tips). Scaphal hooks 
strongly hooked, with a distinct shoulder. Cotor: In life: Colorless, 
transparent, with reddish-orange internal organs, red branchiae, 
golden paleae. 

New records —Arctic ALASKA: Eluitkak Pass, Elson Lagoon near 
Point Barrow, stony; Point Barrow base, washed ashore; off Point 
Barrow base, up to 12.1 miles from shore, 3.7—-123.5 fms., on bottoms 
of mud, stones, mass of worm tubes, and various combinations of 
mud, sand, gravel, rocks, stones, large perforated rocks, shells (17 
stations, 53 specimens); 10 miles west Point Franklin, 13.5 fms., 
sand, Point Barrow Expedition, 1883. Kamcnarxa: Petropavlovsk, 
Grebintzky, 1883. Brrine Sra: Atka Island, Aleutians, Turner, 1879; 
Bering Straits, 13 fms., Dall, 1880. AtasKa: Chichagof Harbor, Attu 
Island, 5-7 fms., 1873; Coal Harbor, Unga Island, 1872; and Port 
Etches, 12-18 fms., 1874; all collected by Dall. Kodiak, Fisher; 
Wrangel, Jones. Wasuincron: Puget Sound, 39-83 fms., mud, rocks, 
shells, Pettibone. Canapian Arctic: Kneeland Bay, Frobisher 
Bay, Baffin Island, and off Daniels Island, Newell Sound, Frobisher 
Bay, 10-30 fms., Bartlett, 1942. Norrawest GREENLAND: Off 
Conical Rock, 76° N., 67°30’ W., 20-40 fms., 1938; between Parker 
Snow Bay and Conical Rock, 25-45 fms., 1940; west side Wolstemholm 
Island, 12 fms., 1940; all collected by Bartlett. East Coast Norra 
America: Off Labrador, 6-12 fms., Blue Dolphin Expeditions, 1949, 
1950, 1951; off New Brunswick, Nova Scotia, Maine, New Hampshire, 
Massachusetts, Long Island Sound, 5-190 fms., U. S. Fish Commission. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Novaya Zemlya. Also 


MARINE POLYCHAETE WORMS—PETTIBONE 313 


Ficure 35.—Sternaspidae: a, Sternaspis scutata, ventral ‘view anterior end; 5, same, ventral 
view posterior end. Pectinariidae: c, Pectinaria hyperborea, ventral view anterior end; 
d, same, dorsal view anterior end; ¢, same, palea; f, same," pectiniform uncinus; g, same, 
dorsal view posterior anal plaque or scapha; h, same, scaphal seta; 7, Pectinaria granulata, 
cephalic paleae; 7, same, cephalic paleae, with short mucronate tips; k, same, scaphal 
hook. (For explanation of symbols, see p. 210.) 


314 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Iceland, Faroes; Labrador to Long Island Sound; Bering Sea to western 
México; north Japan Sea. In low water to 190 fathoms. 


Pectinaria (Cistenides) hyperborea (Malmgren, 1865) 


FiaurReE 35, c—h 


Cistenides hyperborea Malmgren, 1865, p. 360, pl. 18, fig. 40.— Moore, 1903, p. 479.— 
Fauvel, 1914, p. 277, pl. 26, figs. 27, 28.—Augener, 1928, p. 775.—Berkeley 
and Berkeley, 1942, p. 201.—Wesenberg-Lund, 1950a, p. 46; 1950b, p. 106; 
1951, p. 100. 

Pectinaria (Cistenides) hyperborea Hessle, 1917, p. 76.—Nilsson, 1928, p. 31, fig. 
9.—Gustafson, 1936, p. 8.—Okuda, 1937b, p. 56, fig. 5; pl. 2, fig. F—Annen- 
kova, 1937, p. 186; 1938, p. 197.—Zatsepin, 1948, p. 147, pl. 37, fig. 4. 

Description.—Body 28-55 mm. long, 6-12 mm. wide. Tube 45-72 
mm. long, 7-10 mm. in greatest diameter, formed of fine sand grains 
plus some coarser ones. Paleae usually 11-14 pairs (9-15), long, 
tapered to slender tips. Scaphal hooks taper gradually, without 
distinct shoulder. 

New records—Arctic Auaska: Off Point Barrow base, up to 15 
miles from shore, 6-123.5 fms., on bottoms of mud, mass of worm 
tubes, and various combinations of mud, gravel, stones, rocks, with 
shells, worm tubes; from fish trap (8 stations, 19 specimens). Brrine 
Sza: Albatross Sta. 3610, 55°58’ N., 167°16’ W., 75 fms., mud, 1895. 
AuaskKa: Albatross Sta. 4244, Kasaan Bay, Prince of Wales Island, 
50-54 fms. Wurst GREENLAND: Godhavyn, U.S.S. Alert, 1884. East 
Coast Norta America: Off Labrador, 8-125 fms., Blue Dolphin 
Expeditions, 1949, 1950, 1951; off Nova Scotia, Maine, Massachusetts, 
16-65 fms., U. S. Fish Commission. 

Distribution.—Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen, Novaya Zemlya, Kara Sea. 
Also Iceland, Norway to Danish waters, North Sea; Labrador to 
Massachusetts; Bering Sea to Alaska; north Japan Sea to Japan. 
In 1.5-379 fathoms. 


Family AMPHARETIDAE 


Prostomium (or tentacular membrane) more or less distinctly 
trilobed, with numerous filiform oral tentacles, smooth or pinnate, 
retractile in mouth (fig. 36, a, 6, e). Branchiae filiform or subulate 
(rarely pinnate), two to four pairs, inserted on dorsal part of first 
setigerous segments. First few segments achaetous, with or without 
special group of setae or paleae anterior to branchiae. Body divided 
into two distinct regions: (1) thoracic; conical notopodia with smooth, 
limbate capillary setae and, beginning on setigers 3 or 4, with neuro- 
podial flattened pinnules bearing a row of pectiniform uncini; (2) 
abdominal; uncinigerous pinnules only. With or without dorsal, 


MARINE POLYCHAETE WORMS—PETTIBONE 315 


ventral or anal cirri. Tube membranous, covered with mud or ag- 
glutinated foreign bodies. 

Represented by two genera and four species at Point Barrow. 
Both genera have prostomia trilobed, without projecting glandular 
crests; oral tentacles numerous, pinnate; without large dorsal hooks 
posterior to branchiae; branchiae four pairs, filiform or subulate, 
arising on or near a distinct transverse dorsal ridge; thoracic setigers 
14; uncinigerous pinnules begin on setiger 3; notopodia without 
claviform cirri. 


Key to the genera of Ampharetidae from Point Barrow 


1. With paleae, lateral group of golden setae anterior to branchiae (fig. 36, 
eR oe eh I i ee lS ot Ampharete (p. 315) 
Wathout, paledess-. 0 -.6- =25 4952550 ees Asabellides (p. 318) 


Genus Ampharete Malmgren, 1865 


All three species have prostomium distinctly trilobed; median lobe 
with pair of basal eyes, lateral lobes encircling median lobe posteriorly 
(fig. 36, a). First two segments achaetous; first or buccal segment 
extended anteroventrally forming large lower lip. Segment 3 or 
paleal segment with group of paleae arranged in semicircle just ante- 
rior and lateral to branchiae. Abdominal segments with neuropodial 
cirri dorsally on uncinigerous pinnules poorly developed (short, 
rounded to conical) to well developed (cirriform, longer than pinnule). 


Key to the species of Ampharete from Point Barrow 


1. First 2 abdominal segments with enlarged, padlike, achaetous notopodial 
fees eee 2 eS A OD ern Seti Nie ga A. vega 
Without enlarged notopodial lobes on first few abdominal segments----_--_- ») 

2. Paleae slender, taper gradually. Abdominal setigers 12, usually with promi- 
nent neuropodial cirri. Anal cirri numerous (fig. 36, c)_----- A. acutifrons 
Paleae taper rather abruptly, with short mucronate tips. Abdominal setigers 
16-17, with neuropodial cirri inconspicuous. Anal cirri 2___-.-__-_ A. goési 


Ampharete vega (Wirén, 1883) 
FIGuRE 36, a 


Amphicteis vega Wirén, 1883, p. 415, pl. 32, figs. 3, 4. 
Ampharete vega Hessle, 1917, p. 99.—Augener, 1928, p. 778.—Annenkova, 1929, 
p. 493.—Zatsepin, 1948, p. 150. 

Deseription.—Length 16 mm., width 1.5 mm. (up to 50 mm. long, 
4 mm. wide—Wirén, 1883). Paleae slender, tapering gradually to 
fine tips, 10-13 in each group. Eight branchiae form continuous 
transverse group, arising from low fold. Abdominal segments up to 
27-28 (16 in Point Barrow specimen; complete?). First two abdom- 


261112—54——_8 


316 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


inal segments with enlarged, rounded, padlike, achaetous notopodial 
lobes. Neuropodial cirri poorly developed or lacking. Pygidium with 
several small papillae? Colorless in alcohol. Tube thin, membra- 
nous, with sand and debris, rather straggly. 

New records.—Arctic AuAsKA: West side Elson Lagoon near en- 
trance to small lagoon to west, near Point Barrow (rather brackish 
water), 1.2 fms. (1 station, 1 specimen). 

Distribution.—Scattered records in the Arctic: Siberian and Alaskan 
Arctic, Spitsbergen, Barents Sea, Kara Sea, Laptev Sea. In 1.2-11 
fathoms. 


Ampharete acutifrons (Grube, 1860) 
FicureE 36, b-d 


Amphicteis acutifrons Grube, 1860, p. 109, pl. 5, fig. 6. 

Ampharete grubet Malmgren, 1865, p. 363, pl. 19, fig. 44.—Eliason, 1920, p. 70.— 
Fauvel, 1927, p. 227, fig. 79, a-p.—Augener, 1928, p. 776.—Thorson, 1946, 
p. 121, fig. 66—Hartman, 1948, p. 43. 

Ampharete cirrata Webster and Benedict, 1887, p. 747, pl. 8, figs. 110-112. 

Ampharete trilobata Webster and Benedict, 1887, p. 747. 

Ampharete acutifrons Hessle, 1917, p. 96.—Annenkova, 1937, p. 188; 1938, p. 
200.—Zatsepin, 1948, p. 150, pl. 37, fig. 8—Wesenberg-Lund, 1950a, p. 47; 
1950b, p. 109; 1951, p. 102. 


Description.—Length 45 mm., width 6 mm. (up to 56 mm. long, 
8 mm. wide—Wirén, 1883). Paleae slender, taper gradually to long, 
delicate tips, may be somewhat curved, 10-30 in each group. Bran- 
chiae in two groups separated middorsally, each group with three 
branchiae in transverse line and one slightly more posterior. Abdom- 
inal segments 12. With neuropodial cirri small, conical to quite 
long, longer than pinnule (well developed on Point Barrow specimens). 
Pygidium a short ring bearing a circle of numerous subequal anal 
cirri (2 lateral ones may be longer). Colorless in alcohol. Tube 
membranous, covered with soft, gray debris (or agglutinated mud). 

Remarks.—The types of Ampharete cirrata Webster and Benedict 
from Eastport, Maine, were examined. This species was separated 
on the basis of the relative development of the neuropodial cirri, 
which appears to be somewhat variable. The types of A. trilobata 
Webster and Benedict, also from Eastport, were not available, but 
this species also, according to the original description, was differen- 
tiated by the relative development of the cirri. 

New records—Anrctic AtasKa: Eluitkak Pass, Elson Lagoon near 
Point Barrow; Point Barrow base, washed ashore; off Point Barrow 
base, up to 6 miles from shore, 3.7-49 fms., on bottoms of mud, and 
various combinations of mud, pebbles, gravel, stones, rocks, and 


MARINE POLYCHAETE WORMS—PETTIBONE 317 


shells (10 stations, 17 specimens). Wrst Coast Norra AMERICA: 
Washington and Puget Sounds, 80 fms., Pettibone. East Coast 
Norta America: Off Labrador, 8 fms., Blue Dolphin Expedition, 
1949. 

Distribution —Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Jan Mayen, Spitsbergen, Novaya Zem- 
lya. Also Iceland, Swedish west coast to France, Mediterranean; 
Labrador to Maine; Bering Sea to southern California; north Japan 
Sea. In 1-1,333 fathoms. 


Ampharete goési Malmgren, 1865 


Ampharete goést Malmgren, 1865, p. 364, pl. 19, fig. 45.—Hessle, 1917, p. 91.— 
Augener, 1928, p. 778.—Annenkova, 1929, p. 492, pl. 38, fig. 37; 1937, p. 188; 
1938, p. 200.—Zatsepin, 1948, p. 150, pl. 38, fig. 10.—Wesenberg-Lund, 
1950a, p. 47; 1950b, p. 110; 1951, p. 102.— Berkeley and Berkeley, 1952, p. 
66, fig. 136. 

Ampharete goést subsp. braznikovi Annenkova, 1929, p. 492, pl. 38, fig. 44. 

Description Length 35 mm., width 6 mm. (up to 50 mm. long, 7 
mm. wide—Malmgren, 1865). Paleae rather stout, taper abruptly, 
ending in short acuminate tips, 14-21 in each group. Branchiae in 
two groups separated middorsally, each group with three branchiae 
in transverse line. Abdominal segments 17 (stem form and Point 
Barrow specimens) or 16 (subsp. braznikow and Washington Sound 
specimens). Neuropodial cirri inconspicuous, low, rounded. Pygid- 
ium with two long anal cirri. Cotor: In life: Body reddish orange, 
branchiae green. In alcohol: Colorless. 'Tusr: Membranous, rather 
thick, with debris including bits of shells, sea urchins tests, and 
foraminiferans, very stragegly. 

New records—Arctic Ataska: Off Point Barrow base, up to 15 
miles from shore, 20.3-123.5 fms., on bottoms of mass of worm tubes 
and various combinations of mud, pebbles, gravel, stones, rocks, shells, 
and worm tubes (11 stations, 20 specimens). Wersr Coast Nortu 
America: Washington Sound, Pettibone. 

Disiribution—Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen, Barents Sea, White Sea, 
Novaya Zemlya. Also Iceland; British Columbia to Washington; 
Okhotsk Sea to north Japan Sea. In 10.5-353 fathoms. 


Ampharete sp. (Young) 


Two very small specimens, 12 mm. long, were found on bottom of 
rocks with bryozoans and hydroids in 17 fathoms September 9, 1948. 
Prostomium 3-lobed, with four eye spots, bases of four pairs branchiae, 
14 thoracic segments, 10 abdominal segments plus a growing zone. 


318 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Genus Asabellides Annenkova, 1929 


Includes Pseudosabellides Berkeley and Berkeley, 1943. 
Asabellides sibirica (Wirén, 1883) 


FIGuRE 36, e 


Sabellides sibirica Wirén, 1883, p. 418. 

Asabellides orientalis Annenkova, 1929, p. 494, pl. 38, figs. 50, 51, pl. 39, figs. 
60-65; 1937, p. 188. 

Asabellides sibirica Annenkova, 1938, p. 201. 

Pseudosabellides lineata Berkeley and Berkeley, 1943, p. 131; 1944, p. 3; 1952, 
p. 71, fig. 147. 


Description Body long, slender, 38 mm. long, 1.5 mm. wide. 
Prostomium trilobed with median lobe wider and flared anteriorly, 
lateral lobes diagonal. Segments 1 and 2 fused laterally and ventrally, 
forming large lower lip; dorsally, appear as two short segments more 
or less hidden by dorsal branchial fold. Segments 3 and 4 more or less 
fused ventrally and laterally; dorsally, segment 3 without paleae but 
with branchiae; segment 4 with first notopodial lobes and, middorsally, 
a fused nephridial area with pair of papillae posterior to branchial 
bases. Branchiae slender, filiform, emerge from prominent ridge, six 
in transverse row, two slightly posterior. Abdominal segments 23 
(14-23). Neuropodial cirri small, short, rounded (Point Barrow 
specimens) to long, slender, cirriform. Pygidium with few small cirri 
and pair of long lateral ones. Cotor: In life: Salmon below, darker 
above, branchiae green, oral tentacles pink. In alcohol: Colorless. 

New records —Arctic AuasKa: Off Point Barrow base, 5 miles from 
shore, 11.7 fms., on bottom of mud (1 station, 3 specimens). 

Distribution —Scattered records in the Arctic: Siberian, Alaskan, 
and Canadian Arctic. Also Hudson Bay; Bering Sea to British 
Columbia; Okhotsk Sea to north Japan Sea. In 4-106 fathoms. 


Family TEREBELLIDAE 


Body divided into two regions: (1) thoracic; more or less inflated, 
with dorsal bundles of capillary setae and ventral uncinigerous tori 
(sometimes acicular crotchets, sometimes absent), often with ventral 
glandular shields; (2) abdominal; elongated, more or less tapering, 
most often lacking dorsal setae and bearing only uncinigerous pinnules. 
Prostomium of variable form and dimensions, fused with buccal seg- 
ment, forming a lower lip, a transverse dorsal cephalic ridge with or 
without numerous eye-spots, a tentacular membrane with numerous 
filiform grooved tentacles not retractile in mouth, a semicircular or 
folded upper lip (fig. 36, f). Branchiae 1-3 pairs, filiform or arbores- 
cent, or single, quadrilobed, pectinate, on anterior segments, or 


MARINE POLYCHAETE WORMS—PETTIBONE 319 


Ficure 36.—Ampharetidae: a, Ampharete vega, dorsal view anterior end; b, Ampharete acuti- 
frons, dorsal view anterior end; c, same, lateral view posterior end; d, same, palea; ¢, 
Asabellides sibirica, dorsal view anterior end. Terebellidae: f, lateral view anterior end 
of terebellid; g, Amphitrite cirrata, lateral view anterior end; h, same, notoseta; 1, Nicolea 
venustula, lateral view anterior end of male; j, same, thoracic uncini; k, Pista maculata, 
lateral view anterior end, tentacles missing; /, same, uncinus with long manubrium. (For 
explanation of symbols, see p. 210.) 


320 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


branchiae lacking. Without paleae. Tube usually membranous, 
encrusted with sand, debris of shells, algae, etc. 


Represented by 10 genera and 11 species at Point Barrow. 


Key to the genera of Terebellidae from Point Barrow 


Uncini in single rows on first 6 uncinigerous segments, then in 2 rows (alter- 
nating, intercogging or opposite) on certain number of following segments. 
With distinct thoracic ventral shields (single, not paired). Notosetae begin 
on segment 4; uncini begin on segments 5 or 6 (setigers 2 or 3). Fused 
prostomium and buccal segment forming a prominent rectangular lower lip, 
a transverse semicircular cephalic ridge (with or without numerous eye-spots 
along basal groove), flared semicircular upper lip, semicircular depressed 
tentacular region between upper lip and cephalic ridge with numerous, 


filiform, grooved tentacles (fig. 36, f).-=.-...-..- 22 tL Eee 2 
Uncini in single rows only (or with crotchets on thoracic segments)._-.---- 7 

. With 1-3 pairs branchiae dorsally on segments 2-4 (fig. 36, g, 7, k). Uncini 
begin on segment 5 (setiger 2) _.0...:.--.2- 22212 iL Loe 3 
Without branchiae. With lateral lobes on anterior segments (fig. 37, a) ----- 5 


. Notosetae with tips finely denticled (fig. 36, h). Cephalic ridge usually 


without eye-spots. With lateral lobes on anterior segments (fig. 36, g). 
Amphitrite (p. 321) 

Notosetae with tips smooth. Cephalic ridge usually with eye-spots__-_---- 4 
Without lateral lobes on anterior segments. Uncini of anterior segments 
without long manubrium (fig. 36, 7). Branchiae 2 pairs, branched dichoto- 
a sly a (8-36. 4) ee 8 MN ee Nicolea (p.. 322) 
With large lateral lobes on buccal segment, enclosing prostomium and con- 
nected ventrally (fig. 36, k). Uncini of first few segments with long manu- 
brium (fig. 36, 2). Branchiae 1-3 pairs, arborescent, with main trunk well 


markers ei et WO Fe ee ek Pe Ab See Pista (p. 323) 

. Uncini begin on segment 6 (setiger 3). Notosetae of 2 kinds, smooth and 
Gentieled 5 ee ee eu a o_o Proclea (p. 325) 
Uncini begin on segment 5 (setiger 2). Notosetae with tips smooth______- 6 


. Uncini in single rows on first 6 uncinigerous segments, in double rows on next 


10. Nephridial papillae of segment 3 not elongated. Notosetae broadly 
bilimbate below whiplike tip (fig. 37, b)_-.-.------_----- Leaena (p. 325) 
Uncini in single rows on first 6 uncinigerous segments, in double rows on next 
8. Nephridial papillae of segment 3 (anterior to first setiger) elongated. 
Notosetae weakly limbate on one side along most of its length (fig. 37, c). 
Lanassa (p. 326) 


.. Uneini-all short, avicular (ig. 37, d).....-_-22 4... \ ee 8 


Uncini of 2 types; thoracic aciculiform, with long manubrium (fig. 37, A); 
abdominal avicular (fig. 37, 7). Without ventral shields___.__.__.------- 9 


. Branchiae 2-3 pairs, cirriform, in transverse rows. Notosetae begin on 


segment 3, continuing on large number of segments; uncini begin on segment 
5 (setiger 3). Ventral shields usually slightly distinct. Tentacular lobe 
Not especially enlarved = Sn ee Se Thelepus (p. 327) 
Without branchiae. Notosetae begin on segments 2 or 3; uncini begin on 
variable number of segments, may be absent from thoracic region. With 
ventral shields, first unpaired, rest paired, more or less separated in median 
line (fig. 37, f). Tentacular membrane large, trilobed or undulating, 
bearing very numerous tentacles (fig. 37, e-f)---------- Polycirrus (p. 328) 


MARINE POLYCHAETE WORMS—PETTIBONE 321 


9. Branchiae 2-3 pairs, single, filiform (fig. 37, g). Notosetae and uncini 
(crotchets) both begin on segment 6. Tentacular lobe folded, with very 
numerous thin, cylindrical, and short, grooved tentacles. 

Trichobranchus (p. 329) 

Branchiae single, consisting of large cylindrical trunk and 4 pectinate lobes 
(fig. 37, 7). Notosetae begin on segment 3, uncini begin on segment 8 
(setiger 6). Tentacular lobe large, folded, with numerous tentacles of a 
paraded, sees SIA Ba ee) sey Terebellides (p. 330) 


Genus Amphitrite O. F. Miiller, 1771 


Both species have the body inflated anteriorly, decidedly atten- 
uated posteriorly. Cephalic ridge without eye-spots. Branchiae 
three pairs, on segments 2-4. Notosetae begin on segment 4; uncini 
begin on segment 5, in single rows on first six, in double rows on rest 
of thoracic setigers, in single rows on abdominal segments on project- 
ing pinnules. Pygidium a crenulate ring. 


Key to the species of Amphitrite from Point Barrow 


1. Branchiae cirriform, a tuft of simple filaments from short, slightly swollen 
common base (fig. 36, g). Thoracic setigers 17_.____________--- A. cirrata 
Branchiae branched. Thoracic setigers 19_.__._______-___- A. groenlandica 


Amphitrite cirrata Miller, 1776 


Figure 36, g, h 


Amphitrite cirrata O. F. Miller, 1776, p. 216.—Malmgren, 1865, p. 375, pl. 21, 
fig. 53—Webster and Benedict, 1887, p. 748.—Hessle, 1917, p. 185.— 
Chamberlin, 1920, p. 22.—Eliason, 1920, p. 74.—Fauvel, 1927, p. 251, 
fig. 86, i-o.—Augener, 1928, p. 785.—Okuda, 1938b, p. 102.—Hartman, 
1944a, pp. 334, 342; 1948, p. 43.—Thorson, 1946, p. 127, figs. 71, 72.— 
Gorbunov, 1946, p. 39.—Zatsepin, 1948, p. 156, pl. 38, fig. 12.—Wesenberg- 
Lund, 1950a, p. 50; 1950b, p. 116; 1951, p. 106.—Berkeley and Berkeley, 
1952, p. 86, fig. 175. 

Amphitrite radiata Moore, 1908, p. 350; 1923, p. 193. 


Description —Length up to 200 mm., width 10 mm. About 12 
ventral shields. Branchiae 3 pairs, cirriform, a tuft of simple fila- 
ments from a short common base. Each of three branchial segments 
(segments 2-4) with pair of lateral lobes. Nephridial papillae 7 
pairs; on segment 3, prominent papillae ventral to second branchial 
pair; on segments 6-11, rather inconspicuous papillae between noto- 
podia and neuropodia. Thoracic setigers 17. Cotor: In alcohol: 
Reddish brown. Tube: Mostly of mud, with scattered small pebbles 
and debris, soft, breaking easily. 

New records—Arctic AtAsKA: Eluitkak Pass, Elson Lagoon; off 
Point Barrow base, up to 7 miles from shore, 18.3-70 fms., on various 
combinations of mud, sand, gravel, stones, rocks, large perforated 
rocks, shells (8 stations, 14 specimens). Wrst Coast NortH 


322 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


America: Wrangel, Alaska, Jones; Puget Sound, low tide, Pettibone. 
West GREENLAND: Vaigat, Disko Island, Bartlett, 1937. East 
Coast Norra America:| Bay of Fundy, Maine, Massachusetts, 
14-110:fms., U. S. Fish Commission. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitzbergen, Franz Josef Land, 
White Sea, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Nor- 
way to British Isles, Azores, Mediterranean, Adriatic; Bay of Fundy 
to Massachusetts; Bering Sea to southern California; Japan. In low 
water to 1,528 fathoms. 


Amphitrite groenlandica Malmgren, 1865 


Amphitrite groenlandica Malmgren, 1865, p. 376, pl. 21, fig. 52.—Verrill, 1881, 
pp. 305, 310.—Hessle, 1917, p. 181.—Fauvel, 1927, p. 250, fig. 86, a-c.— 
Augener, 1928, p. 787.—Annenkova, 1937, p. 192; 1938, p. 206.—Gorbunov, 
1946, p. 39.—Zatsepin, 1948, p. 157, pl. 38, fig. 14.—Wesenberg-Lund, 
1950a, p. 51; 1950b, p. 117; 1951, p. 107. 

Neoamphitrite robusta Hartman, 1948, p. 44 (not Amphitrite robusta Johnson, 
1901). 

Description.—Length up to 190 mm., width 9 mm. (a single, much 
smaller specimen from Point Barrow). About 14 ventral shields. 
Branchiae 3 pairs, branched dichotomously 3-4 times, with main 
trunk short, thick. Lateral lobes of first two branchial segments well 
developed, continuing ventrally; small lateral lobes on third branchial 
segment. Nephridial papillae 12 pairs on segments 3-14, the first 
four with more prominent papillae. Thoracic setigers 19. Colorless 
in alcohol. ‘Tube of mud. 

Remarks.—A. robusta Johnson is very close to A. groenlandica; 
it differs in having 17 thoracic setigers instead of 19. 

New records —Arctic AutasKA: Off Point Barrow base, 5 miles from 
shore, 49 fms., on bottom of rocks, stones, gravel (1 station, 1 speci- 
men). SouTHWESTERN ALASKA: Round Island, Coal Harbor, Unga 
Island, 8-9 fms., Dall, 1872. 

Distribution.—Scattered records in the Arctic: Siberian and Alaskan 
Arctic, West Greenland, Spitsbergen, Novaya Zemlya. Also Iceland, 
Scandinavian coast to North Sea, Ireland; Maine; southwestern 
Alaska; Okhotsk Sea to north Japan Sea. In 7 to 440 fathoms. 


Genus Nicolea Malmgren, 1865 
Nicolea venustula (Montagu, 1818) 
Figure 36, 1,7 


Terebella venustula Montagu, 1818, p. 344, pl. 13, fig. 2. 

Nicolea arctica Malmgren, 1865, p. 381, pl. 24, figs. 66, 67. 

Nicolea zostericola Malmgren, 1865, p. 381, pl. 26, fig. 76.—Webster and Benedict, 
1887, p. 749.—Moore, 1909b, p. 141.—Fauvel, 1927, p. 261, fig. 90, g—n.— 
Annenkova, 1934, p. 322; 1937, p. 191; 1938, p. 205.—Gustafson, 1936, p. 


MARINE POLYCHAETE WORMS—PETTIBONE aoe 


9.—Berkeley and Berkeley, 1943, p. 180; 1952, p. 87, figs. 177, 178.—Gor- 
bunov, 1946, p. 39.—Hartman, 1948, p. 44. 

Nicolea simplex Verrill, 1873, p. 613; 1881, p. 302, pl. 10, fig. 1—Sumner, Osburn, 
and Cole, 1913, p. 627. 

Nicolea venustula Ehlers, 1913, p. 559.—Hessle, 1917, p. 171—Chamberlin, 1920, 
p. 22.—Eliason, 1920, p. 73.—Fauvel, 1927, p. 260, fig. 90, a-f; 1934a, p. 68.— 
Augener, 1928, p. 788.—Thorson, 1946, p. 126.—Zatsepin, 1948, p. 156, pl. 
38, fig. 11—Wesenberg-Lund, 1950a, p. 51; 1950b, p. 118; 1951, p. 109. 

Description.—Length 15-48 mm., width 2-5 mm., segments 30-46 
(up to 70 mm. long—Wirén, 1883). Ventral shields about 14. 
Cephalic ridge with numerous dark eye-spots. Branchiae two pairs, 
on segments 2-3, branched dichotomously 3-6 times, with very short 
main stem, first pair much larger than second. Without lateral lobes 
on branchial segments. Nephridial papillae on segments 3 (slightly 
posterior to pair of second branchiae), 6, and 7; small in female; in 
male, papillae on segments 6 and 7 long, cylindrical. Thoracic seti- 
gers 15 (14-18, according to Hessle). Uncini begin on segment 5 
(setigerous segment 2), in single rows on first six, in two alternating 
rows on rest of thoracic segments, in single rows on prominent pro- 
jecting abdominal pinnules. Pygidium crenulate. Colorless or 
slightly brownish in alcohol. Females with large yolky eggs inside 
body (Point Barrow, August 21, and September 15, 1948; August 8, 
1949). Tube with tough, translucent membranous lining, with small 
pebbles of various sizes, shell debris, foraminiferans, etc. 

New records—Arctic AtasKa: Off Point Barrow base, up to 12.1 
miles from shore, 13.3-123.5 fms., on bottoms of mud, stones, mass of 
worm tubes, and various combinations of mud, sand, gravel, pebbles, 
stones, rocks, large perforated rocks, with bryozoans, hydroids, shells 
(24 stations, 79 specimens). ALasKa: Security Bay, Jones. Cana- 
p1aAN Arctic: Dobbin Bay, E. Ellesmere Island, 79°36’ N., 73°35’ 
W., 17 fms., Littlewood, 1950. East Coast Norra America: Off 
Maine, Massachusetts, Rhode Island, Connecticut, shore to 63 
fathoms and surface (young), U. S. Fish Commission. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Jan Mayen, Spitsbergen, Franz 
Josef Land, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Nor- 
way to France, Mediterranean, Adriatic, Red Sea; Hudson Bay to 
Connecticut; Bering Sea to western Canada; north Japan Sea; South 
Africa. In low water to 472 fathoms. 


Genus Pista Malmgren, 1865 
Pista maculata (Dalyell, 1853) 


Figure 36, k, l 
Terebella maculata Dalyell, 1853, p. 203, pl. 28, figs. 10-14, 19. 
Scione lobata Malmgren, 1865, p. 383, pl. 23, fig. 62.—Verrill, 1881, pp. 305, 310.— 
Webster and Benedict, 1887, p. 749.—Augener, 1928, p. 789.—Friedrich, 
1939, p. 127. 


324 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Pista maculata Hessle, 1917, p. 161, pl. 3, fig. 4-—Fauvel, 1927, p. 263, fig. 91, 
a-h.—Berkeley and Berkeley, 1943, p. 130.—Gorbunov, 1946, p. 39.— 
Zatsepin, 1948, p. 155, pl. 38, fig. 9—Wesenberg-Lund, 1950a, p. 52; 1950b, 
poit9; 1951,.p.. 110, 

Pista groenlandica Treadwell, 1937, p. 33, figs. 14-16. 


Description —Length up to 150 mm., width 6 mm. Ventral shields 
about 14. Cephalic ridge with numerous eye-spots. Single pair 
large branchiae on segment 2, with large, cylindrical main trunk, 
branched. Buccal segment with two large, rounded lateral lobes 
embracing prostomium, connected ventrally by a crest; segment 2 
short, without lateral lobes; segment 3 with large, rounded, flattened 
lateral lobes. Nephridial papillae on segments 6-7. Thoracic 
setigers 16 (one had small lobe on one side only on 17). Uncini 
begin on segment 5 (setiger 2), in single rows on first 6, in two alternat- 
ing intercogging rows on rest of thoracic segments, in single rows on 
rectangular pinnules in abdominal region. Pygidium with six or 
seven (6-12) conical papillae, arranged starlike. Conor: In alcohol: 
Branchiae and tentacles dark brown or colorless. Tusss: Irregularly 
twisted, with thin transparent membranous lining covered mostly 
with fine sand grains plus few large pebbles, bits of shell, bryozoans, 
foraminiferans, algae, parts of other worm tubes. 

Remarks.—The type of P. groenlandica Treadwell from Baffin Bay 
was examined and is herein referred to P. maculata. Contrary to the 
original description, eye-spots are present on the cephalic ridge (small, 
in transverse groove, easily overlooked). 

New records—Arctic AtasKa: Off Point Barrow base, up to 15 
miles from shore, 21-123.5 fms., on bottoms of mass of worm tubes 
and various combinations of mud, gravel, stones, rocks, large per- 
forated rocks; from crab, Hyas coarctatus (13 stations, 168 plus speci- 
mens). NortH GREENLAND: 5 miles south Cape Chalon; north 
Omenolu near North Star Bay, 17 fms., Bartlett, 1932. Wersr 
GREENLAND: Oelrichs Bay, 1937; off Conical Rock, 76°3’ N., 67°30’ 
W., 20-40 fms., 1938; 1 mile northwest Conical Rock, 25-60 fms., 
1940; between north shores Parkers Snow Bay and Conical Rock, 
25-45 fms., 1940; all collected by Bartlett. East GreEnianp: Off 
Cape Hold with Hope, 23-40 fms., Bartlett, 1939. Easr Coast 
Nort America: Off Labrador, 8-125 fms., Blue Dolphin Expeditions, 
1949, 1950, 1951. 

DMstribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Baffin Bay, Greenland, Spitsbergen, Barent 
Sea, Novaya Zemlya. Also Iceland, Norway to English Channel: 
Hudson Bay to Maine; Bering Sea. In 3-1,528 fathoms. 


MARINE POLYCHAETE WORMS—PETTIBONE 325 


Genus Proclea Saint-Joseph, 1894 


Proclea graffii (Langerhans, 1884) 


Leaena graffiti Langerhans, 1884, p. 262, pl. 15, fig. 21. 

Proclea graffi Southern, 1914, p. 120.—Hessle, 1917, p. 199, fig. 53.— Fauvel, 
1927, p. 268, fig. 94, a-g.—-Gorbunov, 1946, p. 39.—Zatsepin, 1948, p. 157, pl. 
38, fig. 19 —Wesenberg-Lund, 1951, p. 111. 


Description—Length up to 42 mm., width 3 mm., segments 49. 
Ventral shields about 10. Cephalic ridge without eye-spots. With- 
out branchiae. With lateral lobes on segments 2-4, those on segments 
2 and 3 elongate, extending to ventral shields. Nephridial papillae 
on segments 3, 6-8. Thoracic setigers 16. Uncini begin on seg- 
ment 6 (setiger 3), in single rows on first six, in double rows on rest 
of thoracic segments, in single rows on projecting abdominal pinnules. 
Pygidium crenulate. Colorless in alcohol. Tube? 

New records—Arctic AuasKa: Off Point Barrow base, up to 7.5 
miles from shore, 21-36 fms., on bottoms of rocks, stones, large per- 
forated rocks, from breaking rocks apart (3 stations, 6 specimens). 

Distribution.—Scattered records in the Arctic: Siberian and Alaskan 
Arctic, Franz Josef Land, White Sea. Also Bering Sea to Okhotsk 
Sea; Iceland, Swedish west coast, Finland, Ireland, Madeira. In 
1-36 fathoms. 

Genus Leaena Malmgren, 1865 


Leaena abranchiata Malmgren, 1865 


FicureE 37, a, b 


Leaena abranchiata Malmgren, 1865, p. 385, pl. 24, fig. 64.—not Moore, 1909b, 
p. 141 (=Lanassa venusta).—Ehlers, 1913, p. 563.—Hessle, 1917, p. 197.— 
Augener, 1928, p. 793.—Annenkova, 1937, p. 192; 1938, p. 208.—Berkeley 
and Berkeley, 1943, p. 130.—Gorbunov, 1946, p. 39.—Zatsepin, 1948, p. 157, 
pl. 38, fig. 17—Hartman, 1948, p. 45.—Wesenberg-Lund, 1950a, p. 53; 
1950b, p. 121; 1951, p. 112. 

Leaena antarctica McIntosh, 1885, p. 462, pl. 48, figs. 9, 10; pl. 28A, figs. 10, 11. 

Leaena abranchiata var. antarctica Hessle, 1917, p. 197.—Monro, 1930, p. 188; 
1936, p. 178. 


Description.—Length 43 mm., width 3 mm. (up to 75 mm. long, 6 
mm. wide—Malmgren, 1865). About 10 ventral shields. Cephalic 
ridge without eye-spots. Without branchiae. Lateral lobes on seg- 
ment 2 extend ventrally, those on segment 3 extend dorsally, forming 
a prominent ridge (segment preceding first setiger; ridge not so prom- 
inent in var. antarctica). Thoracic setigers 10. Uncini begin on seg- 
ment 5 (setiger 2), in single rows on first 6, in double rows on next 10, 
then in single rows on prominent abdominal pinnules. Notosetae 
widely bilimbate below whiplike tips. Colorless in alcohol. TusE: 
Of soft mud with a few bits of scattered rock (some small ones on 
tubes of Pista maculata). 


326 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 108 


New records —Arctic AtasKka: Off Point Barrow base, 25 fms., on 
bottoms of gravel, stones, shells (1 station, 1 specimen). East Coast 
Nortsa America: Off Labrador, 15-55 fms., mud and mud with rock, 
Blue Dolphin Expeditions, 1950, 1951. 

Distribution.—Widely distributed in the Arctic: Siberian and Alas- 
kan Arctic, Baffin Bay, Greenland, Spitsbergen, Novaya Zemlya, 
Kara Sea. Also Iceland, Faroes, Norway, Sweden, Finland; Hudson 
Bay to Labrador; southwestern Alaska; Okhotsk Sea to north Japan 
Sea. In 6-833 fathoms. Var. antarctica: Antarctic, South Georgia; 
up to 1,975 fathoms. 


Genus Lanassa Malmgren, 1865 
Lanassa venusta (Malm, 1874) 
FIGURE 37, c 


Laphaniella venusta Malm, 1874, p. 98, pl. 1, fig. 8. 
Leaena nuda Moore, 1905b, p. 855, pl. 44, figs. 14, 15. 

Leaena abranchiata Moore, 1909b, p. 141 (not L. abranchiata Malmgren, 1865). 
Lanassa venusta Hessle, 1917, p. 205.—Eliason, 1920, p. 75.—Gustafson, 1936, 
p. 9.—Annenkova, 1937, p. 194; 1938, p. 209.—Zatsepin, 1948, p. 157. 

Lanassa venusta subsp. pacifica Annenkova, 1938, pp. 209, 230. 

Description.—Length 37-55 mm., width 3 mm., segments 54-67. 
Ventral shields about 10. Cephalic ridge without eye-spots (present in 
subsp. pacijica). Oral tentacles numerous (more than 12 as indicated 
by Malm). Without branchiae. Lateral lobes on segments 2 and 3 
extending ventrally; small lateral lobes on segment 4. Nephridial 
papillae on segments 3, 6-8; those of segment 3 prominent (anterior 
to first setiger). Thoracic setigers 11 (12 on subsp. pacifica). Uneini 
begin on segment 5 (setiger 2), single on first six, in double rows on 
next eight, in single rows on prominent abdominal pinnules. Noto- 
setae weakly limbate (fig. 37, c). Pygidium crenulate. Females 
with large yolky eggs in body (Point Barrow, Sept. 9 and 15, 1948; 
Aug. 17, Oct. 11, 1949; Aug. 1, 1950). Conor: In life: Orange, with 
white shields, tentacles tan. In alcohol: Colorless. Tus: With 
thin, transparent membranous lining, with very loose mud, bits of 
sand, rock, foraminiferans (tube of specimen from Labrador with 
great deal of loosely assembled shell debris). 

Remarks.—The type of Leaena nuda Moore from southwestern 
Alaska was examined and found in poor condition. It has been referred 
previously to Lanassa venusta by Annenkova (1937). 

New records—Arctic ALASKA: Point Barrow base, washed ashore; 
off Point Barrow base, up to 12.1 miles from shore, 18.3—75.5 fms., on 
bottoms of mud, stones, and various combinations of mud, sand, 
gravel, pebbles, stones, rocks, large perforated rocks, with bryozoans, 
shells (17 stations, 44 specimens). CaNnapian Arctic: Cove in 


MARINE POLYCHAETE WORMS—PETTIBONE 327 


Kneeland Bay, Frobisher Bay, Baffin Island, 14 fms., Bartlett, 1942. 
Distribution Scattered records in the Arctic: Siberian, Alaskan, 
and Canadian Arctic. Also Swedish west coast, Danish waters; 
Labrador; southwestern Alaska; north Japan Sea. In 7-140.5 
fathoms. 
Genus Thelepus Leuckart, 1849 


Thelepus cincinnatus (Fabricius, 1780) 
Fiagure 37, d 


Amphitrite cincinnata Fabricius, 1780, p. 286. 

Thelepus circinnata Malmgren, 1865, p. 387, pl. 22, fig. 58. 

Thelepus cincinnatus Webster and Benedict, 1887, p. 749.—Hessle, 1917, p. 212.— 
Chamberlin, 1920, p. 23.—Eliason, 1920, p. 76.—Fauvel, 1927, p. 271, fig. 
95, i-m; 1932, p. 233, fig. 40; 1934a, p. 69.—Augener, 1928, p. 790.—Monro, 
1930, p. 192; 1936, p. 182.—Treadwell, 1937, p. 33.—Friedrich, 1939, p. 127.— 
Hartman, 1944a, pp. 336, 343; 1952, p. 236.—Gorbunov, 1946, p. 39.— 
Zatsepin, 1948, p. 154, pl. 38, fig. 7.—Wesenberg-Lund, 1950a, p. 54; 
1950b, p. 122; 1951, p. 112. 

Thelepus hamatus Moore, 1905b, p. 856, pl. 44, figs. 16-18.—Hartman, 1948, 
p. 44.—Berkeley and Berkeley, 1952, p. 82, fig. 167. 

Description.—Up to 200 mm. long, 10 mm. wide, segments nu- 
merous. Dorsal surface rugose, with small glandular warts irregularly 
distributed and especially abundant anteriorly. Ventral shields in- 
distinct, wide, somewhat wrinkled. Buccal segment and prostomium 
forming a thick lower lip, a prominent horseshoe-shaped upper lip, 
a semicircular cephalic ridge with numerous small eye-spots; tentacular 
area between upper lip and cephalic ridge with numerous, rather 
thick tentacles. Branchiae 2 pairs on segments 2 and 3, each formed 
of numerous, simple filaments arranged in parallel transverse rows. 
Nephridial papillae on segments 4-7. Notosetae begin on segment 3, 
continuing on about 40 segments (or almost to posterior end). Un- 
cini begin on segment 5 (setiger 3), in single rows throughout, the 
uncinigerous tori gradually transformed into projecting rectangular 
pinnules. Pygidium with crenulate anal opening. Conor: In life: 
Orange with red branchiae, tan tentacles, with whitish glandular 
areas ventrally and laterally. In alcohol: Colorless. Tus: Cylin- 
drical, twisted, with tough, transparent, membranous lining, covered 
with small pebbles of varying sizes, with fragments of shells, bryo- 
zoans, foraminiferans, old worm tubes, algae. 

Remarks.—The type of T. hamatus Moore from Alaska was ex- 
amined and is herein referred to 7. cincinnatus; it is a small specimen, 
consisting of anterior end only. 

New records—Axrctic Auaska: Off Point Barrow base, up to 15 
miles from shore, 20—123.5 fms., on bottoms of mass of worm tubes, 
rocks, stones, gravel, attached to stems of Tubularia (6 stations, 7 
specimens). Wurst GREENLAND: Godhavn, off Hare Island, 70°20’ 


328 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


N., 56° W., 90 fms., Greely Relief Expedition, 1884. One mile 
northwest Conical Rock, 25-60 fms., and north shore Wolstenholm, 
13-25 fms., Bartlett, 1940. East Greentanp: Off Cape Hold with 
Hope, 23-40 fms., Bartlett, 1939. East Coast Norta America: 
Off Labrador, 12-30 fms., Blue Dolphin Expedition, 1949; off New 
Brunswick, Nova Scotia, Maine, New Hampshire, Massachusetts, 16- 
640 fms., U. S. Fish Commission. Wersr Coast Norra America: 
Washington and Puget Sounds, low water to 46 fms., mud, shell, rock, 
Pettibone. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Jan Mayen, Spitsbergen, Franz 
Josef Land, Barents Sea, Novaya Zemlya. Also Iceland, Faroes, 
Shetlands, Norway to Portugal, Azores, Madeira, Mediterranean, 
Adriatic; Labrador to Massachusetts; Bering Sea to Washington; 
Japan; Indian Ocean (Andamans). Southern latitudes: South Geor- 
gia, South Orkneys, South Shetlands, Graham Coast, Palmer 
Peninsula. In low water to 1,391 fathoms. 


Genus Polycirrus Grube, 1851 
Polycirrus medusa Grube, 1855 
Figure 37, e, f 


Polycirrus medusa Grube, 1855, p. 120.—Hessle, 1917, p. 220.—Fauvel, 1927, p. 
279, fig. 97, a~-d.—Augener, 1928, p. 795.—Annenkova, 1934, p. 322; 1937, 
p. 194; 1938, p. 210.—Gorbunov, 1946, p. 39.—Zatsepin, 1948, p. 154, pl. 38, 
fig. 5—Wesenberg-Lund, 1950a, p. 54; 1950b, p. 123; 1951, p. 114. 

Ereutho smitti Malmgren, 1865, p. 391, pl. 23, fig. 63.—Webster and Benedict, 
1887, p. 749. 

Polycirrus sp., Hartman, 1948, p. 45. 

Description—Length up to 70 mm., width 6 mm. Integument 
tesselated. Large unpaired ventral shield on first few segments, 
followed by about six pairs of ventral shields separated in midline. 
Prostomium and buccal segment forming lower lip, wide semicircular 
dorsal cephalic ridge (without eye-spots), large trilobed or undulating 
tentacular membrane with very numerous tentacles on its outer edge, 
the inner part forming folded upper lip; some tentacles long, slender, 
filiform, others thicker, distinctly grooved. Thoracic setigers usually 
13 (10-13), beginning on segment 3 (or 2). Without thoracic uncini; 
uncini begin on segment 16 (or 15 or just posterior to last setiger), 
uniserial, few in number, very small and inconspicuous, on slightly 
raised abdominal pinnules. Six pairs nephridial papillae on segments 
3-8. Females with large, yolky eggs (Point Barrow, September 15, 
1948, and October 14, 1949). Conor: In life: Red. In alcohol: 
Colorless. Tus: Simple galleries in mud or sand. 

New records—Arctic AtasKa: Off Point Barrow base, up to 15 
miles from shore, 13.3-78.2 fms., on bottoms of mud and various com- 


MARINE POLYCHAETE WORMS—PETTIBONE 329 


binations of mud, pebbles, gravel, stones, rocks, large perforated 
rocks, with bryozoans, worm tubes (13 stations, 35 specimens). 
SPITSBERGEN: Spitsbergen Sea, U.S. S. Alliance, 1881. Nortrawexst 
GREENLAND: One mile northwest Conical Rock, 25-60 fms., Bartlett, 
1940. Brrine Sea: St. Paul Island, Pribilofs, Palmer, 1890. East 
Coast Norra America: Off Labrador, 45 fms., silt, Blue Dolphin 
Expedition, 1949. Wrst Coast Norra America: Pavlof Bay, 
Alaska, 150 fms., Alaska King Crab Investigation, 1940; Washington 
Sound, 12-46 fms., mud and mussels, Pettibone. 
Distribution.—Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen, White Sea, Novaya Zemlya, 
Kara Sea. Also Iceland, Faroes, Swedish west coast to France, 
Mediterranean; Labrador to Maine; Bering Sea to Washington; 
Okhotsk Sea to north Japan Sea. In low water to 889 fathoms. 


Genus Trichobranchus Malmgren, 1865 
Trichobranchus glacialis Malmgren, 1865 
FIcurRE 37, g-1 


Trichobranchus glacialis Malmgren, 1865, p. 395, pl. 24, fig. 65.—Webster and 
Benedict, 1887, p. 750.—Ehlers, 1913, p. 566.—Hessle, 1917, p. 131.— 
Fauvel, 1927, p. 288, fig.100,a-h.—Augener, 1928, p. 792.—Annenkova, 
1937, p. 190; 1938, p. 202.—Zatsepin, 1948, p. 153, pl. 38, fig. 1—Wesenberg- 
Lund, 1950a, p. 55; 1950b, p. 125; 1951, p. 115.—Berkeley and Berkeley, 
1952, p. 76, figs. 154, 155. 

Trichobranchus glacialis var. antarcticus Hessle, 1917, p. 132.—Hartman, 1952, 
p. 233. 


Description.—Length up to 30 mm., width 3 mm., segments 60-70. 
Buccal segment and prostomium form thick, tesselated, inflated lower 
lip connected laterally with pair of projecting, rounded, flattened lobes 
(ventral to mass of tentacles), cephalic ridge with numerous black 
eyespots (absent in var. antarcticus), undulating tentacular membrane 
with very numerous tentacles, and folded upper lip; some tentacles 
larger, distinctly grooved; others smaller, filiform. Three pairs 
branchiae on segments 2-4, each composed of single long filament. 
With slightly projecting lateral lobes on branchial segments. Thoracic 
setigers and uncinigers 15, beginning on segment 6; notosetae limbate, 
with smooth capillary tips; neuropodial uncini aciculiform (crotchets). 
Abdominal segments with avicular uncini in single rows on triangular 
projecting pinnules. Pygidium with anal opening crenulate. Color- 
less in alcohol. Tube membranous, encrusted with mud or fine sand. 

New records—Arctic Autaska: Off Point Barrow base, up to 5 
miles from shore, 27-49 fms., on various combinations of mud, gravel, 
stones, rocks, shells (5 stations, 15 specimens). East Coast Nort 
America: Cape Cod Bay, Massachusetts, 27-118 fms., U. S. Fish 
Commission. 


330 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Distribution.—Widely distributed in the Arctic: Alaskan Arctic, 
Greenland, Spitsbergen, Novaya Zemlya, Kara Sea. Also Iceland, 
Faroes, Swedish west coast to Spain, Canary Islands, Mediterranean; 
Maine to Massachusetts; British Columbia; north Japan Sea. Var. 
antarcticus: Magellan Straits, South Georgia, Ross Island, Kaiser 
Wilhelm II Land. In 3-1,517 fathoms. 


Genus Terebellides M. Sars, 1835 
Terebellides stroemii M. Sars, 1835 
Fiaure 37, j-m 


Terebellides stroemit M. Sars, 1835, p. 48, pl. 13, fig. 31—Malmgren, 1865, p. 
396, pl. 20, fig. 48.—Webster and Benedict, 1887, p. 750.—Moore, 1903, p. 
478; 1908, p. 352; 1923, p. 199.—Hessle, 1917, p. 137—Chamberlin, 1920, 
p. 23.—Eliason, 1920, p. 72.—Fauvel, 1927, p. 291, fig. 100, i-q; 1932, p. 
234; 1947, p. 79, fig. 76, f-i—Augener, 1928, p. 797.—Gustafson, 1936, p. 
9.—Annenkova, 1937, p. 190; 1938, p. 202.—Treadwell, 1937, p. 35.—Okuda, 
1938b, p. 102.—Friedrich, 1939, p. 127.—Berkeley and Berkeley, 1943, p. 
130; 1944, p. 5; 1952, p. 75, figs. 152, 153.—Hartman, 1944a, pp. 336, 343; 
1944d, p. 24; 1951, p. 113.—Thorson, 1946, p. 124, fig. 69.—Gorbunov, 1946, 
p. 39.—Zatsepin, 1948, p. 153, pl. 38, fig. 3—St¢gp-Bowitz, 1948c, p. 68.— 
Wesenberg-Lund, 1949, p. 355; 1950a, p. 55; 1950b, p. 126; 1951, p. 115, fig. 
11.—Hartman and Reish, 1950, p. 44. 

Terebellides stroemii var. japonica Moore, 1903, p. 478; 1908, p. 352; 1923, p. 200. 

Description.—Length up to 75 mm., width 8 mm., segments 50-60. 

Prostomium and buccal segment forming large, transverse, crescent- 

shaped plate below mouth, large and undulating tentacular membrane 

with numerous, short, grooved tentacles, folded upper lip; dorsally 
without cephalic ridge or eye-spots. Segment 2 short, achaetous. 

Single branchia formed of large cylindrical trunk dorsally on segment 

3 (first setiger) and four pectinate lobes, frequently interlocked giving 

appearance of more or less solid mass. With somewhat developed 

lateral lobes extending ventrally on segments 2—7 (larger on segments 

3-6). Thoracic setigers 18, begin on segment 3. Uncini begin on 

segment 8 (setiger 6), those on first unciniger consisting of crotchets 

with long manubrium and bent, pointed tips, those on rest of thoracic 
segments long, aciculiform, with large fang and few denticles; may be 
crowded in more than one row in large specimens. Abdominal 
uncini short, avicular, pectiniform, in single rows on projecting tri- 

angular pinnules. Prominent paired nephridial papillae on segment 3, 

smaller ones on segments 6 and 7. Pygidium with terminal anus, with 

sides crenulate. Female with large yolky eggs (ripe eggs and sperm, 

September 26, 1949). Conor: In life: Body orange, branchia orange- 

red, tentacles tan. In alcohol: Colorless, with smooth iridescent 

surface. 
Parasites——One of the 87 specimens had the parasitic copepod 

Saccopsis terebellidis Levinsen (identified by P. L. Illg) attached to 


MARINE POLYCHAETE WORMS—PETTIBONE 331 


Ficure 37.—Terebellidae: a, Leaena abranchiata, lateral view anterior end; », same, notoseta; 
¢, Lanassa venusta, notoseta; d, Thelepus cincinnatus, avicular uncinus; e, Polycirrus 
medusa, dorsal view anterior end; f, same, ventral view anterior end; g, Trichobranchus 
glacialis, dorsal view anterior end; h, same, thoracic aciculiform uncinus; i, same, abdom- 
inal avicular uncinus; 7, Terebellides stroemii, lateral view anterior end; k, same, crotchet 
from first unciniger; /, same, thoracic aciculiform uncinus; m, same, abdominal avicular 
uncinus. (For explanation of symbols, see p. 210.) 


261112—54——_9 


382 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


anterior part in the region of the branchia. The same polychaete and 
copepod species were found in Iceland waters and figured by Wesen- 
berg-Lund (1951, p. 117, fig. 11). 

New records.—Arctic AtasKa: West side Elson Lagoon near Point 
Barrow, 1.2 fms.; Eluitkak Pass, Elson Lagoon; Point Barrow base, 
washed ashore; off Point Barrow base, up to 15 miles from shore, 18.3— 
123.5 fms., on bottoms of mud, stones, mass of worm tubes, and various 
combinations of mud, sand, pebbles, gravel, stones, rocks, large per- 
forated rocks, with shells, bryozoans, worm tubes (26 stations, 87 
specimens). Wrst Coast Norta America: Washington and Puget 
Sounds, 10-165 fms., Pettibone. East Coast Norra America: Off 
Labrador, 5-65 fms., Blue Dolphin Expeditions, 1949, 1950; off Nova 
Scotia, Maine, Massachusetts, Rhode Island, Long Island Sound, 
6-368 fms., U. S. Fish Commission. 

Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Jan Mayen, Spitsbergen, Franz 
Josef Land, Barents Sea, White Sea, Novaya Zemlya, Kara Sea. Also 
Iceland, Faroes, Norway to Portugal, Mediterranean, Adriatic, Black 
Sea; Hudson Bay to Long Island Sound, Gulf of Mexico, West Indies; 
Bering Sea to southern California, Panamaé, Venezuela; north Japan 
Sea to Japan; Iranian Gulf; Indian Ocean; South Africa, Bouvet 
Island, Kerguelen, New Caledonia. In low water to 1,611 fathoms. 


Family SABELLIDAE 


Body cylindrical, tapered posteriorly. Prostomium indistinct. 
Buccal region with mouth terminal, with dorsal lip, two ventral lips 
or vesicular bulbs, two fleshy, membranous or filiform palps of variable 
length. Branchiae form conspicuous, often highly colored terminal 
funnellike plume surrounding mouth; plume formed of two semi- 
circular or spiral lobes bearing numerous filaments each with two rows 
of ciliated barbules (fig. 38, a). Branchiae without operculum. A 
collarette more or less developed on first segment. Body divided 
into two regions: (1) thoracic; few segments (4-12), with dorsal bun- 
dles of capillary setae and ventral uncinigerous tori (tori lacking in 
Myzicola) ; without thoracic membrane; (2) abdominal; with inversion 


Ficure 38.—Sabellidae: a, Sabella crassicornis, lateral view anterior end; b, same, tip 
of branchial filament; c, same, thoracic notosetae; d, same, dorsal view pygidium; ¢, same, 
thoracic hoe-like seta from ventral torus; f, same, thoracic avicular uncinus from ventral 
torus; g, same, ventral view collarette; 4, same, lateral view; i, same, dorsal view; 7, Pot- 
amilla neglecta, dorsal view collarette; k, same, lateral view; /, same, ventral view; m, 
same, dorsal thoracic limbate seta; m, same, dorsal thoracic spatulate setae; 0, Potamilla 
reniformis, ventral view collarette; p, same, lateral view; g, same, dorsal view; 7, same, 
portion of branchial filament; s, same, thoracic avicular uncinus from ventral torus; #, 
same, thoracic hoe-like seta from ventral torus; u, same, tip of tube, rolled at free end. 
(For explanation of symbols, see p. 210.) 


333 


MARINE POLYCHAETE WORMS—PETTIBONE 


ee facing page. 


Figure 38.—For explanation s 


334 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


of setae, dorsal uncinigerous tori and ventral capillary setae. A 
median ciliated faecal groove runs length of body, ventral in abdom- 
inal region and crossing to dorsal surface in thoracic region. Tube 
cylindrical, permanent or more or less transitory, gelatinous, mem- 
branous, or horny, covered or not with mud, sand, gravel, debris of 
shells. 

Represented by five genera and seven species at Point Barrow. 
All the genera have large branchial plumes composed of two similar 
semicircular lobes; branchial filaments without large subterminal 
eyes, without dorsal appendages. First setigerous segments with 
limbate notosetae only; neurosetae begin on setiger 2. Abdominal 
region with numerous segments. 


Key to the genera of Sabellidae from Point Barrow 


1. Ventral tori of thorax with row of uncini of the avicular form (fig. 38, f; in 
Sabella and Potamilla with additional row of hoelike setae, fig. 38, t). Col- 
larette well developed. Branchial filaments with or without eyes, united 
at base only, with palmar membrane poorly developed or lacking------_- 2 

Ventral tori of thorax with crotchets with long manubrium (fig. 39, #; without 
tori in Myzicola). Branchial filaments without eyes, united along a large 
part of their length by a well-developed palmar membrane (fig. 39, m)_-. 3 

2. Dorsal thoracic setae all limbate, some long with straight borders, others short 

With wide borders.(fig. SS, c)o2- = oe es Sabella (p. 334) 
Dorsal thoracic setae of two kinds, some limbate, others spatulate (fig. 38, m, n) 
Potamilla (p. 335) 

3. Uncinigerous tori projecting, with short, lateral rows of uncini. Collarette 
well developed (fig. 39, a—c). Dorsal thoracic setae of two kinds, limbate 
and spatulate (fig. 39, f, 9). 22-----.--------_-.--2, -2-- 4 

Without projecting uncinigerous tori, abdominal uncini in almost complete 
transverse band (fig. 39, p). Collarette poorly developed, represented by 
a triangular ventral lobe (fig. 39, 0). Dorsal thoracic setae all capillary, 


limbate so 2 2 See ee BA Ae ee ek ee ee Myxicola (p. 340) 

4. Posterior segments without ventral depression--_--..-.------- Chone (p. 337) 
Posterior segments about 10, with ventral suckerlike disc, large ventral de- 
pression with flared sides (fig. 39, n)-.----------------- Euchone (p. 339) 


Genus Sabella Linné, 1767 (sensu Malmgren, 1865) 
Sabella crassicornis Sars, 1851 
FIGuRE 38, a-7 


Sabella crassicornis Sars, 1851, p. 202.—Malmgren, 1865, p. 399, pl. 27, fig. 83.— 
Moore, 1909b, p. 144.—Johansson, 1927, p. 119.—Hartman, 1942a, p. 78; 
1948, p. 46.—Berkeley and Berkeley, 1943, p. 130; 1952, p. 114, figs. 236, 
237.—Zatsepin, 1948, p. 161, pl. 39, fig. 4. 

Sabella fabricit Kroyer, 1856, p. 20.—Fauvel, 1927, p. 300, fig. 103, a-g—_Augener, 
1928, p. 800.—Annenkova, 1934, p. 322; 1937, p. 195; 1938, p. 211.— Wesen- 
berg-Lund, 1950b, p. 128; 1951, p. 118. 

Sabella spetsbergensis Malmgren, 1865, p. 399, pl. 29, fig. 93. 

Sabella spitzbergensis Webster and Benedict, 1887, p. 750. 


MARINE POLYCHAETE WORMS—PETTIBONE 335 


Description.—Length 20-80 mm., width 3-4 mm. Collarette 
widely separated middorsally, with midventral slit and lateral notches, 
resulting in 4-lobed structure; ventral lobes closely approximated, 
may be deflected. Two branchial lobes each with about 16 filaments 
(15-35), with short tapering tips. Branchial filaments with usually 
four to six (2-8) pairs of eyes (located in color bands of filaments and 
may easily be overlooked). Thoracic setigers usually 8 (7-9). Abdo- 
men with one or two pairs of eye-spots between parapodial rami, well 
developed toward posterior end. Pygidium with pair of bulbous 
lobes with groups of eye-spots on dorsolateral sides. Conor: In 
life: Body flesh color, with branchial filaments banded rusty red, 4-7 
transverse bands per filament. In alcohol: Body colorless with red- 
dish purple bands on branchial filaments. Tuss: Cylindrical, free 
end flexible, covered with thin, smooth layer of mud; embedded part 
transparent, horny, rigid, covered with sand grains and foreign 
material. 

New records—Arctic AuAsKA: Off Point Barrow base, up to 15 
miles from shore, 36—123.5 fms., on bottoms of stones, mass of worm 
tubes, and various combinations of gravel, stones, rocks, large per- 
forated rocks, with worm tubes (8 stations, 15 specimens). Kam- 
CHATKA: Petropavlovsk, Grevnitzky, 1888. AtasKa: New Harbor, 
Unga Island, under stones, Dall, 1872; Albatross Sta. 2847, 55°01’ N.., 
160°18’ W., 48 fms., and station at Kodiak, 1888. Canapran Arctic: 
Foxe Basin, 25-31 fms., Bartlett, 1927. East Coast Nort AMERICA: 
Off Labrador, 5-6 fms., Blue Dolphin Expedition, 1949; Bay of Fundy, 
Maine, Massachusetts, 15 fms., U. S. Fish Commission. CENTRAL 
America: Golfo Dulce, west Costa Rica, M. Valerio. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Novaya Zemlya, Kara 
Sea. Also Iceland, Faroes, Norway to France, Mediterranean; 
Hudson Bay to Massachusetts; Bering Sea to California, Central 
America (Costa Rica); north Japan Sea to Japan. In low water to 
230 fathoms. 

Genus Potamilia Malmgren, 1865 


Key to the species of Potamilla from Point Barrow 


Pav thont branchial eyes... ale. eee edness S. oel eee P. neglecta 
With compound eyes in single rows on branchial filaments, 0-8 per filament 
gts [oe Es OEE OE bo beta 2 ET ee ce) teeta ge ee P. reniformis 


Potamilla neglecta (Sars, 1851) 


FicureE 38, j-n 


Sabella neglecta Sars, 1851, p. 203. 
Potamilla neglecta Malmgren, 1865, p. 401, pl. 27, fig. 84.—Webster and Benedict, 
1884, p. 736.—Moore, 1909b, p. 145; 1923, p. 242.— Johansson, 1927, p. 143.— 


336 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Augener, 1928, p. 801.—Hartman, 1942a, p. 81; 1944a, pp. 336, 343; 
1948, p. 46.—Zatsepin, 1948, p. 162, pl. 39, fig. 7—Wesenberg-Lund, 1950a, 
p. 56; 1950b, p. 128; 1951, p. 119.—Berkeley and Berkeley, 1952, p. 116, fig. 
238. 

Potamilla torelli Malmgren, 1865, p. 402; 1867, p. 114, pl. 13, fig. 76.—Ehlers, 
1913, p. 575.—Fauvel, 1927, p. 310, fig. 107, m—-s; 1934a, p. 71.—Annenkova, 
1934, p. 322; 1937, p. 195; 1938, p. 213.—Wesenberg-Lund, 1951, p. 121. 

Potamilla acuminata Moore and Bush, 1904, p. 159, pl. 11, figs. 3-6; pl. 12, fig. 
41. 

Description.—Length 30-84 mm., 1-3 mm. wide. Collarette widely 
separated dorsally, sloping ventrally, with midventral slit, entire 
laterally, resulting in a 2-lobed structure; ventral lobes triangular, 
may be deflected. Branchial lobes each with about 16 filaments 
(6-20), with short, naked distal tips, without eyes. Thoracic setigers 
usually 8 (5-8). Pygidium with pair of bulbous lobes with dark eye- 
spots. Conor: In alcohol: Body colorless, branchiae with two or 
three diffused reddish brown bands. Tuse: Horny, transparent, 
more or less covered with sand or mud; smaller tubes with rather 
uniform layer of small sand grains; larger tubes may be encrusted 
with living barnacles, bryozoans, hydroids, foraminiferans, amphipod 
tubes. Large yolky eggs in a single layer, pressed close to wall, 
about one-third way down in tube; a transparent, thin membrane 
laid down between eggs and worm (large developing yellow eggs 
found in tube, September 6, 1949, dredged in 36 fathoms, Point 
Barrow). 

New records—Arctic ALASKA: Off Point Barrow base, up to 12.1 
miles from shore, 21.7-123.5 fms., on bottoms of mass of worm tubes 
and pebbles, gravel, rocks, large stones (7 stations, 24 specimens). 
Brrine Sea: Albatross Sta. 3496, 56°32’ N., 169°45’ W., 41 fms., 
1893; St. Paul Island, Pribilofs, Hahn, 1911. Anasxa: Albatross 
Sta., Kodiak, 1888. Wurst Greentanp: Off Hare Island, 70°20’ N., 
56° W., 90 fms., U.S. S. Alert, 1884. Easr Coast Norto Ammrica: 
Off Labrador, 125 fms., Blue Dolphin Expedition, 1949; off Maine, 
Massachusetts, 31-130 fms., U. S. Fish Commission. 

Distribution—Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Spitsbergen. Also Iceland, Faroes, Nor- 
way to France, Madeira, Mediterranean, Adriatic, Cape Verde 
Islands; Labrador to Massachusetts; Bering Sea to California; north 
Japan Sea to Japan; Antarctic. In low water to 1,044 fathoms. 


Potamilla reniformis (Leuckart, 1849) 


FIGureE 38, o-u 


Sabella reniformis Leuckart, 1849, p. 183, pl. 3, fig. 8. 
Potamilla reniformis Malmgren, 1867, p. 114, pl. 13, fig. 77—Webster and Bene- 
dict, 1884, p. 736; 1887, p. 750.—Johansson, 1927, p. 142.—Fauvel, 1927, 


MARINE POLYCHAETE WORMS—PETTIBONE sar 


p. 309, fig. 107, a-l._—Okuda, 1937b, p. 61.—Annenkova, 1934, p. 322; 1937, 
p. 195; 1938, p. 212.—Zatsepin, 1948, p. 161, pl. 39, fig. 6 —Wesenberg-Lund, 
1950a, p. 57; 1950b, p. 129; 1951, p. 120. 

Pseudopotamilla intermedia Moore, 1905c, p. 562, pl. 37, figs. 15-22; 1908, p. 359.— 
Hartman, 1938c, p. 25, pl. 2, fig. 8; pl. 3, figs. 1-4; 1948, p. 46.—Rioja, 
1941, p. 732. 

Pseudopotamilla reniformis Hartman, 1944a, pp. 336, 343, pl. 21, figs. 3, 6; 1945, 
p. 47.—Berkeley and Berkeley, 1952, p. 116, fig. 239. 

Description —Length 80-100 mm., width 1-2 mm. Collarette 
with middorsal depression, deeply notched dorsolaterally (at level of 
notopodia), with midventral slit, resulting in a 4-lobed structure of 
pair of rounded or triangular dorsal lobes and pair of rounded ventral 
lobes which may be deflected. Branchial lobes each with about 10 
filaments (5-15). At least some of branchial filaments with large 
compound eyes, usually 0-3 per filament (0-8), eyes may be variable 
in size or subequal, deep reddish in alcohol. Thoracic setigers usually 
10 (7-14). Pygidium with pair of bulbous lobes with eye-spots. 
Cotor: In alcohol: Body colorless, branchiae with diffused reddish 
brown bands in region of eyes. TuBE: Somewhat twisted, horny, 
translucent or opaque, more or less covered with sand or mud, the 
free end usually flattened, rolled scroll-like when animal pulls inside. 

New records—Arctic AuasKa: Off Point Barrow base, up to 7.4 
miles from shore, 21.7-54.6 fms., on bottoms of pebbles, gravel, 
rocks, in holes of large stones (2 stations, 15 specimens). Brrine 
Sea: Anchorage, Cape Etoliu, Nunivak Island, 8 fms., Dall, 1874. 
SouTHEASTERN ALasKA: Lituya Bay, 6-9 fms., Dall, 1874. Easr 
Coast Norro America: Off Nova Scotia, Maine, New Hampshire, 
Massachusetts, Rhode Island, Delaware, 10-317 fms., U. S. Fish 
Commission. 

Distribution Scattered records in the Arctic: Arctic Alaska, 
Greenland. Also Iceland, Swedish west coast to France, Mediter- 
ranean; Nova Scotia to North Carolina; Bering Sea to British Co- 
lumbia, México; north Japan Sea to Japan. In low water to 317 
fathoms. 


Genus Chone Kroyer, 1856 


Both species have the segments biannulate. Collarette deeply 
incised middorsally, forming two longitudinal-folded brackets, with 
sides smooth; without midventral or lateral notches (fig. 39, a-c). 
Branchial filaments united on great part of their length by palmar 
membrane (fig. 39, m). Thoracic setigers 8. Thoracic dorsal setae 
of three kinds—limbate capillary setae, fine bayonette setae, and 
spatulate setae with asymmetrical mucronate tips or gently rounded 
(fig. 39, f-h). Pygidium with anus terminal, with rounded to conical 
bulbous dorsal valve (fig. 39, d). 


338 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Key to the species of Chone from Point Barrow 
1. Collarette nearly straight (fig. 39, b). Free end of branchial filaments flat- 


tened, foliaceous, widely limbate (fig. 39, e).___._.-___- C. infundibuliformis 
Collarette oblique, longer on ventral side (fig. 39, k). Free end of branchial 
filaments with long filiform tips (fig. 39, l)._....._.._.____--____- C. dunéri 


Chone infundibuliformis Kroyer, 1856 
Figure 39, a-j 


Chone infundibuliformis Kréyer, 1856, p. 33.—Malmgren, 1865, p. 404, pl. 28, 
fig. 87.—Théel, 1879, p. 66.—Wirén, 1883, p. 422.—Fauvel, 1927, p. 334, 
fig. 116, a-o.—Augener, 1928, p. 806.—Annenkova, 1934, p. 322; 1937, 
p. 196; 1938, p. 215.—Hartman, 1942b, p. 136; 1944a, pp. 334, 336 (not pl. 
20, fig. 5), pl. 21, fig. 7—Berkeley and Berkeley, 1943, p. 130; 1952, p. 123, 
figs. 252, 253.—Gorbunov, 1946, p. 39.—Zatsepin, 1948, p. 164, pl. 39, 
fig. 13.—Wesenberg-Lund, 1950a, p. 58; 1950b, p. 131; 1951, p. 123. 

Chone gracilis Moore, 1906b, p. 257, pl. 12, figs. 62-66.—Berkeley and Berkeley, 
1952, p. 123, fig. 254. 


Description.—Length 30 mm., width 2.5 mm. (up to 120 mm. long, 
6 mm. wide—Fauvel, 1927). Collarette nearly straight laterally. 
Branchial filaments about 15 (10-36), with tips more or less foliaceous, 
edged by transparent border (prolongations of palmar membrane). 
Cotor: In life: Variable—flesh color, branchiae orange spotted with 
white; olive green with distal half of branchiae brick red, basal half 
red and olive, chalk white on outer sides of bases of branchiae. In 
alcohol: Colorless, grayish, or tannish. Tuss: Membranous, en- 
crusted with sand, mud, or pebbles of variable sizes, foraminiferans. 

New records—Arctic ALASKA: Hluitkak Pass, Elson Lagoon near 
Point Barrow, 6.6 fms.; off Point Barrow base, up to 7.5 miles from 
shore, 20-49 fms., on bottoms of mud, stones, and various combina- 
tions of mud, sand, gravel, stones, rocks, large perforated rocks, shells 
(12 stations, 32 specimens). Canapran Arctic: East side Cobourg 
Island, Baffin Bay, 75° 40’ N., 78° 40’ W., Bartlett, 1935. NorrH- 
west GREENLAND: 1 mile northwest Conical Rock, Bartlett, 1940. 
East GREENLAND: Off Cape Hold with Hope, 23-40 fms., Bartlett, 
1939. SprrsperGEen: Spitsbergen Sea, U.S.S. Alliance, 1881. Brrine 
Sea: St. Paul Island, Pribilofs, Palmer, 1890; St. George Island, 
Pribilofs, Hanna, 1913; Albatross Sta. 3232, 58° 30’ N., 157° 34’ W., 
10.5 fms., 1890, Sta. 3233, 58° 23’ N., 157° 42’ W., 7 fms., 1890, and 
Sta. 3289, 56° 44’ N., 159° 16’ W., 16 fms., 1890; Kiska, Aleutians, 
Dall. Wesr Coast Nortu America: Strait of Juan de Fuca, Wash- 
ington, 40 fms., mud, Pettibone. East Coast Nortu America: Off 
Nova Scotia, Maine, Massachusetts, Rhode Island, 10-206 fms., U. 
S. Fish Commission. 

Distribution—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Novaya Zemlya, Kara 


MARINE POLYCHAETE WORMS—PETTIBONE 339 


Sea. Also Iceland, Faroes, Shetlands, Scandinavian coast to Danish 
waters; Hudson Bay to Rhode Island; Bering Sea to California; north 
Japan Sea. In low water to 1,955 fathoms. 


Chone dunéri Malmgren, 1867 
Ficure 39, k, | 


Chone dunért Malmgren, 1867, p. 116, pl. 13, fig. 75.—Wirén, 1883, p. 422.— 
Fauvel, 1927, p. 336, fig. 117, l-r; 1934a, p. 74.—Augener, 1928, p. 807.— 
Monro, 1936, p. 189.—Treadwell, 1937, p. 35.—Berkeley and Berkeley, 1944, 
p. 5.—Zatsepin, 1948, p. 164, pl. 39, fig. 15.—Wesenberg-Lund, 1950a, p. 
58; 1950b, p. 130; 1951, p. 123.—Hartman, 1951, p. 117. 

Description —Length 20-35 mm., width 1.5-2.5 mm. Collarette 
oblique, longer on ventral side. Branchial filaments about 10 (5-11), 
with long, tapered tips. Co tor: In life: Branchiae yellow. In alco- 
hol: Colorless or tannish. Tusse: Thin, membranous, encrusted with 
sand. 

New records —Arctic AuasKa: Off Point Barrow base, up to 7.5 
miles from shore, 13.3-49 fms., on bottoms of mud and various com- 
binations of mud, sand, gravel, stones, rocks, large perforated rocks, 
shells (9 stations, 21 specimens). CaNnapr1AN Arctic: Baffin Island, 
66° 43’ N., 80° 07’ W., Bartlett, 1927. Easr Coast Nort AmMmRrIca: 
Off Labrador, 8 fms., mud, Blue Dolphin Expedition, 1949. 

Distribution —Widely distributed in the Arctic: Alaskan and Ca- 
nadian Arctic, Greenland, Spitsbergen, Novaya Zemlya, Kara Sea. 
Also Iceland, Faroes, Norway to North Sea, Madeira, Mediterranean, 
Adriatic; Labrador; Florida; west coast South America (Pert). In 
8-889 fathoms. 


Genus Euchone Malmgren, 1865 
Euchone analis (Kroyer, 1856) 
FIGURE 39, m, n 


Sabella analis Kréyer, 1856, p. 17. 

Euchone analis Malmgren, 1865, p. 406, pl. 28, fig. 88.—Théel, 1879, p. 65.— 
Wirén, 1883, p. 423.—Chamberlin, 1920, p. 27.—Augener, 1928, p. 804.— 
Annenkova, 1937, p. 196; 1938, p. 216.—Berkeley and Berkeley, 1943, p. 
130; 1952, p. 121, figs. 250, 251.—Gorbunov, 1946, p. 39.—Zatsepin, 1948, 
p. 163, pl. 39, fig. 12—Wesenberg-Lund, 1950a, p. 59; 1950b, p. 182; 1951, 
p. 124. 

Description —Length 19-28 mm., width 1.8 mm. (up to 60 mm. 
long—Augener, 1928). Segments biannulate and, with the ventral 
faecal groove, forming four large areas per segment. Collarette 
with opening dorsally, nearly straight and entire laterally, with 
shallow midventral incision. Branchial filaments about 10 per lobe 
(up to 14), united through a great part of their length by a well- 
developed palmar membrane. Thoracic setigers 8. Ventral anal 


340 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


depression extending over 10-12 most posterior segments, bordered 
by high sinuous membrane. Pygidium with terminal anus, with 
conical bulbous dorsal valve. Colorless or tannish in alcohol. Tube 
membranous, encrusted with sand grains and certain amount of 
debris. 

New records.—Arctic AtasKa: Off Point Barrow base, up to 12.1 
miles from shore, 36—123.5 fms., on bottoms of mass of worm tubes 
and stones, rocks, large perforated rocks (3 stations, 3 specimens). 
East Coast Norto America: Off Labrador, 30 fms., mud with 
rock, Blue Dolphin Expedition, 1950. 

Distribution —Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Davis Strait, Greenland, Jan Mayen, Spits- 
bergen, Novaya Zemlya, Kara Sea. Also Iceland, Norway to 
Danish waters; Hudson Bay to Labrador; Bering Sea to British 
Columbia; north Japan Sea. In 2-389 fathoms. 


Genus Myxicola (Koch in MS.) Rénier, 1848 
Myxicola infundibulum (Montagu, 1808) 
Figure 39, o-q 


Amphitrite infundibulum Montagu, 1808, p. 109, pl. 8. 

Myzicola steenstrupii Kréyer, 1856, p. 35.—Malmgren, 1865, p. 408, pl. 29, fig. 
90.—Théel, 1879, p. 66.—Webster and Benedict, 1884, p. 737; 1887, p. 
750.—Moore, 1909b, p. 144.—Eliason, 1920, p. 79.—Zatsepin, 1948, p. 164, 
pl. 39, fig. 16. 

Myzxicola infundibulum Fauvel, 1927, p. 342, fig. 119, a-i; 1934a, p. 74; 1936, 
p. 86.—Annenkova, 1934, p. 322.—Okuda, 1939, p. 243.—Hartman, 1944a, 
pp. 335, 343, pl. 21, fig. 1; 1948, p. 47.—Wesenberg-Lund, 1950b, p. 
134; 1951, p. 125.—Berkeley and Berkeley, 1952, p. 119, fig. 244. 


Description —Length 23 mm., width 5 mm. (up to 200 mm. long, 
15 mm. wide—Fauvel, 1927). Collarette represented by a triangular 
ventral lobe extending anteriorly between the branchial lobes, absent 
laterally and dorsally. A middorsal groove on first 8-10 segments. 
Branchial filaments 20-40, united for greater part of their length by 


Ficure 39.—Sabellidae: a, Chone infundibuliformis, ventral view collarette; b, same, lateral 
view; ¢, same, dorsal view; d, same, dorsal view pygidium; ¢, same, tip of foliaceous bran- 
chial filament; f, same, thoracic dorsal limbate capillary seta; g, same, thoracic dorsal spatu- 
late seta; h, same, thoracic dorsal bayonette seta; 7, same, thoracic crotchet from ventral 
torus; 7, same, abdominal uncinus; k, Chone dunéri, lateral view collarette; J, same, tip 
of branchial filament; m, Euchone analis, lateral view anterior end; n, same, ventral view 
posterior end; 0, Myxicola infundibulum, lateral view anterior end; p, same, lateral view 
segment from abdominal region; g, same, dorsal view pygidium. Serpulidae: r, Spirorbis 
granulatus, dorsal view tube; s, same, operculum (after Fauvel); t, same, collar seta (seta 
of first setiger); u, Spirorbis spirillum, dorsal view anterior end extended from tube (var. 
ascendens, after Emerton); v, same, discoidal form of tube; w, same, operculum; x, same, 
collar seta (seta of first setiger, after Fauvel). (For explanation of symbols, see p. 210.) 


MARINE POLYCHAETE WORMS—PETTIBONE 341 


WY Will 
LA 


Figure 39.—For explanation see facing page. 


342 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


well-developed palmar membrane; tips of filaments limbate. Thora- 
cic setigers 8 (7-9). Thoracic notopodia with limbate capillary setae; 
neuropodia with crotchets with long manubrium (may disappear). 
Abdomen without projecting tori; avicular uncini in an almost com- 
plete circle; with capillary neurosetae. With one to several lateral 
eye-spots behind each parapodia. Pygidium with anus terminal, 
with rounded, bulbous dorsal valve with lateral groups of eye-spots. 
Cotor: In alcohol: Body irregularly speckled reddish brown, with 
branchiae violet distally. Tusr: Gelatinous, transparent, very 
firm and elastic, may be quite thick. 

New records.—Arctic AtasKa: Off Point Barrow base, up to 4.25 
miles from shore, 25-35.5 fms., on bottoms of mud with gravel (2 
stations, 2 specimens). Wrst Coast Norra America: Washington 
and Puget Sounds, low water to 165 fms., Pettibone. Easr Coast 
Norra America: Off Nova Scotia, Maine, Massachusetts, 9-90 
fms., U. S. Fish Commission. 

Distribution —Scattered records in the Arctic: Arctic Alaska, 
West Greenland, Novaya Zemlya. Also Iceland, Finland, to English 
Channel, Mediterranean, Adriatic; Nova Scotia to Massachusetts; 
Bering Sea to California; Japan. In low water to 287 fathoms. 


Family SERPULIDAE 


Body cylindrical or fusiform, slightly flattened. Prostomium indis- 
tinct. Buccal region with mouth terminal, with two transverse lips, 
palps absent or slightly developed. Branchiae form a terminal funnel- 
like plume around mouth, formed of two semicircular or spiralled 
lobes bearing few to numerous filaments, each with two rows of ciliated 
barbules (fig. 39, u). Usually with one or two opercula (calcareous, 
horny, or membranous) formed on the first dorsal filaments of the 
branchial lobes. Usually with a collarette on first segment. Body 
divided into two regions: (1) thoracic; with few segments (3-7), with 
dorsal bundles of capillary setae and ventral uncinigerous tori; usually 
with a thoracic membrane; (2) abdominal; with inversion of setae, 
dorsal uncinigerous tori and ventral capillary setae. A wide, shallow, 
longitudinal ciliated faecal groove, ventral in abdominal region, dorsal 
in thoracic. Tube cylindrical or polygonal, calcareous, opaque or 
rarely transparent, fixed to substratum (rarely free). 

Represented by a single genus and two species (two subgenera) at 
Point Barrow. 


Genus Spirorbis Daudin, 1800 


Both species are of small size, asymmetrical and coiled. Branchial 
lobes each with 3-5 filaments. A single, partly calcareous operculum 


MARINE POLYCHAETE WORMS—PETTIBONE 343 


on smooth peduncle. Collarette large, widely separated dorsally, 
entire laterally and ventrally. Three thoracic setigers, the first 
setiger with only dorsal setae (collar setae) and the next two with 
ventral tori in addition to dorsal setae. With well-developed thoracic 
membrane. A long achaetous region between thorax and abdomen 
(may be crowded with ova; spermatozoa develop in more posterior 
segments; hermaphroditic). About 20 abdominal setigers. Pygidium 
with two rounded lobes. Colorless in alcohol. Tube small (1-3 mm. 
in diameter), closely coiled, white, opaque, fixed to hydroids, bryozoa, 
algae, carapaces of crustaceans, spines of tunicates, stones, etc. 


Key to the species of Spirorbis from Point Barrow 


1. Tube with sinistral (counterclockwise) spiral coiling (with mouth of tube 
facing observer, the opening is on the left; fig. 39, r). Eggs incubated in 
operculum. Collar setae with well-developed basal crenulate wings (fig. 
eI Pree ea Ss ean aS ee ee Say S. (Laeospira) granulatus 

Tube with dextral (clockwise) spiral coiling (fig. 39, v). Eggs incubated in 
tube. Collar setae without crenulate wings (fig. 39, x). 
S. (Dexiospira) spirillum 


Spirorbis (Laeospira) granulatus (Linné, 1767) 
Figure 39, r-t 


Serpula granulata Linné, 1767, p. 1266. 
Spirorbis quadrangularis Stimpson, 1854, p. 29.—Moore, 1908, p. 362.— Hartman, 
1944a, pp. 336, 343. 
Spirorbis (Laeospira) granulatus Borg, 1917, p. 28, figs. 14-16.—Fauvel, 1927, p. 
403, fig. 137, q-u.—Augener, 1928, p. 815.—Annenkova, 1937, p. 198; 1938, 
p. 219.—Berkeley and Berkeley, 1943, p. 130; 1952, p. 137, figs. 286, 287.— 
Gorbunov, 1946, p. 39.—? Thorson, 1946, p. 139, fig. 80.—Zatsepin, 1948, 
p. 166.—Wesenberg-Lund, 1950a, p. 63; 1950b, p. 141; 1951, p. 135. 
Description.—Tube 1-2 mm. in diameter, dull chalky white, opaque, 
somewhat rugose and variable with two longitudinal ridges, one on 
each side (the opening almost quadrangular), or with three more or 
less distinct longitudinal keels, or without crests; the tube is coiled 
sinistrally, up to two coils, flatly on smooth surfaces or somewhat 
open when on rough surfaces; it may encircle a strand of hydroid and 
the free end may extend upward. Operculum with terminal plate 
strongly convex, calcareous, with long, cylindrical projection (short 
on one side, longer on opposite side), with large brood pouch for 
incubating eggs (eggs in operculum, Point Barrow, August 17, 1949, 
and February 18, 1950). Collar setae with blades finely serrated, 
with well-developed crenulate wing at base. 
New records—Arctic Auaska: Off Point Barrow base, up to 12.1 
miles from shore, 27-123.5 fms., on bottoms of mass of worm tubes, 


344 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


and various combinations of mud, gravel, rocks, stones (3 stations, 7 
specimens). SovuTHWESTERN ALAsKA: Kiska Harbor, Aleutians, 10 
fms., Dall, 1873; Canoe Bay, 25 fms., on old clam shell, Alaska King 
Crab Investigation, 1940. East Coast Norta America: Off Lab- 
rador, 30-35 fms., mud, rocks, Blue Dolphin Expedition, 1950; off 
Nova Scotia, Maine, Massachusetts, North Carolina (Cape Hatteras), 
17-34 fms., U. S. Fish Commission. 

Distribution —Widely distributed in the Arctic: Siberian and 
Alaskan Arctic, Greenland, Jan Mayen, Spitsbergen, Franz Josef 
Land, Novaya Zemlya, Kara Sea. Also Iceland, Faroes, Norway to 
France; Hudson Bay to North Carolina; Alaska to British Columbia; 
north Japan Sea. In low water to 239 fathoms. 


Spirorbis (Dexiospira) spirillum (Linné, 1758) 
FIGURE 39, u-« 

Serpula spirillum Linné, 1758, p. 785. 

Spirorbis (Dexiospira) spirillum Borg, 1917, p. 20, figs. 3, 4—Fauvel, 1927, p. 
392, fig. 132, f-p.—Augener, 1928, p. 814.—Annenkova, 1934, p. 322; 1937, 
p. 197; 1938, p. 218.—Treadwell, 1937, p. 35.—Hartman, 1942a, p. 91; 
1944a, pp. 336, 343, pl. 22, fig. 2—Berkeley and Berkeley, 1943, p. 130; 
1952, p. 133, figs. 272-274.—Thorson, 1946, p. 138.—Gorbunov, 1946, p. 
39.—Zatsepin, 1948, p. 166.—Wesenberg-Lund, 1950a, p. 62; 1950b, p. 138; 
1951, p. 132. 

Circeis spirillum Chamberlin, 1920, p. 28. 

Dexiospira spirillum Hartman, 1944b, p. 287; 1948, p. 51; 1951, p. 121. Hartman 
and Reish, 1950, p. 47. 

Description.—Tube 1-3 mm. in diameter, white, opaque (may be 
somewhat translucent), shiny, smooth, porcellaneous, coiled dextrally, 
up to 3% coils; tube may be coiled flatly (discoidal form) when 
attached to smooth surfaces, or partly unrolled, the last coil or part 
of last coil raised from substratum (var. ascendens Levinsen, var. 
lucidus Mérch) when attached to rough surfaces, as branching 
bryozoan colonies. Operculum with shallow, calcareous, concave, 
terminal plate, with slight projection (talon) on under side. Collar 
setae with serrate blades, without basal crenulate wings. Embryos 
incubated in tube (eggs in tube, Point Barrow, September 9, 1948, 
and August 30, 1949). 

New records—Arctic AuasKa: Off Point Barrow base, up to 8 
miles from shore, 21-75.5 fms., on bottoms of stones and various 
combinations of gravel, rocks, on bryozoa, on Hyas coarctatus, on 
spines of Boltenia echinata (8 stations, 39 specimens). GULF OF 
Auaska: Albatross Sta., Observation Island, Cordova, 1914. West 
GREENLAND: Godhavn, Greely Relief Expedition, 1884. East Coast 
Norts America: Off Labrador, 10-40 fms., Blue Dolphin Expeditions, 
1949, 1950, 1951; off Nova Scotia, Maine, Massachusetts, Long 
Island Sound, 16-83 fms., U. S. Fish Commission. 


MARINE POLYCHAETE WORMS—PETTIBONE 345 


Distribution.—Widely distributed in the Arctic: Siberian, Alaskan, 
and Canadian Arctic, Greenland, Spitsbergen, Franz Josef Land, 
Kara Sea. Also Iceland, Faroes, Norway to France; Hudson Bay to 
Long Island Sound, southern Texas; Bering Sea to México; north 
Japan Sea to Japan. In low water to 183 fathoms. 


346 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


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MARINE POLYCHAETE WORMS—PETTIBONE 355 


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356 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


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Vol. 103 Washington : 1954 No. 3325 


THE RELATIONSHIPS OF OLD AND NEW WORLD 
MELANIANS 


By J. P. E. Morrison 


Recent anatomical observations on the reproductive systems of 
certain so-called ‘‘melanian”’ fresh-water snails and their marine rela- 
tives have clarified to a remarkable degree the supergeneric relation- 
ships of these fresh-water forms. 

The family of Melanians, in the broad sense, is a biological ab- 
surdity. We have the anomaly of one fresh-water “family” of snails 
derived from or at least structurally identical in peculiar animal 
characters to and ancestrally related to three separate and distinct 
marine families. On the other hand, the biological picture has been 
previously misunderstood largely because of the concurrent and 
convergent evolution of the three fresh-water groups, Pleuroceridae, 
Melanopsidae, and Thiaridae, from ancestors common to the marine 
families Cerithiidae, Modulidae, and Planaxidae, respectively. 

The family Melanopsidae is definitely known living only in Europe. 
At present, the exact placement of the genus Zemelanopsis living in 
fresh waters of New Zealand is uncertain, since its reproductive 
characters are as yet unknown. In spite of obvious differences in 
shape, the shells of the marine genus Modulus possess at least a well- 
indicated columellar notch of the aperture, to corroborate the biologi- 
cal relationship indicated by the almost identical female egg-laying 
structure in the right side of the foot of Modulus and Melanopsis. 


273553—54——1 357 


358 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


The family Pleuroceridae, fresh-water representative of the ancestral 
cerithiid stock, is now known to include species living in Africa, 
Asia, and the Americas. 

The family Thiaridae, likewise, is known from Africa, Asia, and 
America. With identical reproductive characters, the Thiaridae are 
only too obviously the fresh-water relatives of the Planaxidae. It is 
only when the Melanopsidae, Pleuroceridae, and Thiaridae are 
separated that their zoogeographic story can be read without confu- 
sion. Perfect separation of these three fresh-water families can only 
be accomplished by reference to characters of the animals, particularly 
the reproductive characters. Radular, opercular, or shell characters 
that will completely separate these families are nonexistent because of 
their concurrent and convergent evolution. 

The following key to the major groups of the Melanian complex is 
given here as the most logical one possible. In the author’s opinion, 
it represents the true biological relationship of these groups. 


a.1 Reproduction dioecious; males present in species. 
b.1 Female with large grooved ‘‘ovipositor”’ in pit on right side of foot; few eggs 
of proportionately large size. 
ci. Marine shelled jecees - geht -e 5- reper tendo leieretaree Modulidae 
e2. Wresh-water shells...¢¢ 2 20 $2 Fo RE ee ee Melanopsidae 
b.2 Female with egg-laying sinus and rudimentary papilla, or no special struc- 
ture on right side of foot; numerous eggs of proportionately smaller size. 
d.! Marine shells. .-.._..-..-.......s3.)1¢6 Aw SW=ee7)) oo Cerithiidae 
a2 °Fresh:water:shelisivin besos. 20! of. epeingial tn Pleuroceridae 
e.1 Males with no intromittent structures. 
f Females oviparous, with egg-laying sinus in right side of foot. 


Pleurocerinae 

f2 Females ovoviviparous, without egg-laying sinus; brood pouch 
uterine, the enlarged end of the oviduct_________-_--- Lavigeriinae 

e2 Males with eversible ‘‘penis’”’ in mantle edge near end of vas deferens; 
females ovoviviparous; brood pouch uterine___________- Tiphobiinae 


a.2 Reproduction parthenogenetic; no males present in species; brood pouch not 
uterine, but adventitious (subhaemocoelic) in the neck region, with opening 
on right side of neck. 

g:? *Marine ‘shelissi1 0 4 LULA MiOvI OO) EF 98 D1eG ON RIO as Wee Planaxidae 
g2iFresh=water'shells:. 2 990 Sil} to jerartanalc inpeya ont 30 Thiaridae 


Family MELANOPSIDAE (Europe) 


Two genera from the fresh waters of Europe have been studied and 
proven to belong to the family Melanopsidae. They are more special- 
ized than the Asiatic and American Pleuroceridae, possessing a much 
larger ‘‘ovipositor” and laying a much smaller number of proportion- 
ately much larger eggs. 


OLD AND NEW WORLD MELANIANS—MORRISON 359 


According to studies by Ankel (1928), Fagotia esperi (Ferussac) 
has the same reproductive anatomy of the female as does Melanopsis. 
The few large eggs (1 mm. im size) are laid singly in an irregular capsule. 

Melanopsis dufourit (Ferussac) from Elche, Alicante Province, 
Spain, has been examined by me. Unlike the animals of the North 
American Pleuroceridae, in this species at least, the smooth mantle 
edge is continuous beneath the back of the foot, forming a circle around 
the aperture. In the female the end of the oviduct is quite far behind 
the mantle edge and simple, as is typical for the group. The egg- 
laying groove is not long, and does not even reach the mantle edge. 
The pit is enormously developed and has a large, whitish, tongue- 
shaped, basally-attached ‘‘ovipositor,” with a groove on its posterior 
dorsal face, toward the rest of the pit. According to Ankel (1928), 
a few proportionately large eggs are laid by Melanopsis. 


Family PLEUROCERIDAE (in the Americas) 
Subfamily PLEUROCERINAE 


All known American members of Pleuroceridae are oviparous, 
including those from North, Central, and South America and the 
West Indies. In other words, every American genus and species for 
which the characters of reproduction are known is egg-laying, with an 
egg-laying sinus or pit in the right side of the foot of the females. The 
presence of this pit is the single morphological character that proves 
them to belong to the typical subfamily. There is every indication 
that all the American genera and species of Pleuroceridae belong here, 
although some of the North American genera are still unrecorded as to 
critical animal characters. 


Subfamily PLEUROCERINAE (in North America) 
Genus Oxytrema Rafinesque, 1819 


Ozxytrema Rafinesque is the earliest and correct name for one of the 
most widespread ‘‘Melanian” genera in the world. This genus in- 
cludes numerous North American species whose ranges extend from 
the Atlantic to the Pacific coasts and from southern Canada to Florida 
and Texas. It also includes North American fossils, as well as a num- 
ber of Recent species from southeast Asia (Korea, China, and Thailand). 
All the species called ‘‘Pleurocera” by Bryant Walker, and other 
authors who followed him blindly, and all the species called ‘“‘Gonz- 
obasis” (with very few exceptions) belong to this genus. Their eggs 
are laid in a single row in a close, irregularly spiral group, in apparent 
flat clusters of 3 to 10 egg capsules in each small egg mass, the whole 
covered with sand grains as recorded by Van Cleave (1932), Winsor 
(1933), and Woodward (1934). 


360 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Rafinesque originally described Oxytrema (1819, p. 423), including 
young specimens of both his genus Pleurocera and this one. There 
were no specific names included. Blainville, in 1824 and againin 
1825 (p. 442), placed one species of Rafinesque under this generic 
name. He gave as the sole example of the subgenus Pleurocerus 
(Oxytrema) acutus (Rafinesque), validating the specific name in 1824. 
Rafinesque (1831, p. 3) again described his Pleurocera acuta, and at 
the same time declared he had given the name in 1818. From 1824 
on, Oxytrema has been the earliest available name for the group be- 
cause the genotype was fixed at that time as Pleurocera (Oxytrema) 
acuta Blainville by monotypy. With no serious question ever raised 
about the identity of acuta, the genotype, doubts about the identity 
of the genus Oxytrema vanish. 

The eastern American (Appalachian) species of Oxytrema include 
some whose shells are almost completely smooth as adults, such as 
O. symmetrica (Haldeman); some with spiral sculpture predominant, 
such as O. virginica multilineata (Say); others with axial sculpture 
strong, such as QO. laqueata (Say); and still others with both axial 
and spiral lirae to produce reticulate or nodose sculpture, such as 
O. catenaria (Say). 

In exactly parallel fashion, as should be expected of congeneric 
stocks, the western American species now known to belong to Oxytrema 
show the same rather complete range of sculpture from smooth to 
axially and spirally sculptured adult shells. The easternmost living 
representative of the Rocky Mountain group of species is 0. comalensis 
(Pilsbry). This species from Texas belongs to this minor group 
within the genus, possessing the same minutae of female reproductive 
characters as does O. plicifera (Lea) from Oregon and Washington. 
Most if not all the western America Tertiary fossil species described 
as “Melania,” ‘“Goniobasis,” and ‘‘Pachychilus,”’ from Texas to 
Washington, were members of this genus Ozytrema Rafinesque. For 
example, ‘Amblorus’’ olequaensis Arnold and Hannibal (Hannibal, 
1912, p. 178, pl. 8, fig. 27) closely resembles the smoother phase of 
the living Ozxytrema silicula (Gould) from the same region, while 
“Pachychilus” drakei Arnold and Hannibal (Hannibal, 1912, p. 183, 
pl. 8, fig. 26) is close to the living species Oxytrema plicifera (Lea). 
Certain other fossil species from the Rocky Mountain region parallel 
eastern forms in the possession of cancellate or reticulate sculpture. 
Species known from the fossil record indicate the previous continuity 
of geographic distribution from Washington to Texas, now noncon- 
tinuous since much of the intervening area (progressively desiccated 
since the Miocene) is now unsuited to survival of pieurocerine fresh- 
water snails. 

Female animals of Oxytrema canaliculatum undulatum (Say) from 


OLD AND NEW WORLD MELANIANS—MORRISON 361 


the Kentucky River, of Oxytrema deshayesiana (Lea) from the Tennes- 
see River, of Oxytrema bulbosa (Gould) from eastern Oregon, and of 
Oxyirema nodifila (Martens) from the Han River, Korea, have been 
examined by me and sketched to show the egg-laying groove and the 
“ovipositor” pit on the right side of the foot (see pl. 11). 


Section Strephobasis Lea, 1861 


The group named Strephobasis by Lea (1861, p. 96), with the geno- 
type Oxytrema (Strephobasis) plena Anthony, as stated by Pilsbry 
(1896a, p. 496), is only an extreme section of the genus, according to 
shell characters. This small group, which is confined to the larger 
rivers of the upper Tennessee drainage, is exactly parallel to Goniobasis, 
sensu stricto, from the Coosa River drainage. The animal characters 
must be examined to determine whether this group is identical to or 
biologically distinct from Ozytrema, sensu stricto, and Goniobasis, 
sensu stricto. 

Genus Gyrotoma Shuttleworth, 1845 


Gyrotoma Shuttleworth (1845, p. 88), based on the genotype 
(Gyrotoma ovoideum Shuttleworth=) Gyrotoma excisum (Lea), is con- 
fined to the Coosa River drainage. As Goodrich (1924, p. 6) has 
hinted, the group named Goniobasis by Lea in 1862 (genotype by 
subsequent designation by Hannibal 1912: G. osculata Lea) may 
eventually prove to be most closely related to Gyrotoma. It is possible 
that Goniobasis, sensu stricto, may be a subgenus of Gyrotoma different 
only by lack of or development of the fissure in the lip. Study of the 
animal characters (as yet unknown) of the genotype must prove the 
true biological position of Goniobasis. 


Genus Mudalia Haldeman, 1840 


Mudalia of Haldeman, whose genotype is Mudalia carinata (Bru- 
guiére) from the Atlantic coastal region, includes trilineata (Say) of the 
Ohio River and the more widespread ‘‘Goniobasis’”’ livescens (Menke) 
from the upper St. Lawrence River and upper Mississippi River areas. 
The single carina typical of adolescent shells and the egg-laying habits 
are identical in carinata (Winsor, 1933) and in livescens (Jewell, 1931). 
Figures 10 and 14 of Goodrich (1945, pl. 1) show how closely the shape 
of adult shells of livescens may vary toward the usual shape of adult 
shells of carinata. The eggs of Mudalia are laid singly; that is, there is 
only one egg capsule in each egg mass. 


Genus Leptoxis Rafinesque, 1819 


Leptoxis of Rafinesque is the earliest name available for the Ameri- 
can species related to the genotype L. praerosa (Say), and must be so 
used. Pilsbry (1917, p. 113) has stated the case for priority of Lep- 


362 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


toris Rafinesque 1819 over Anculosa Say 1821 so clearly that Walker’s 
1918 refusal to follow this needed correction of nomenclature appears 
at present to be only a prejudiced personal objection to the change. 
As in other similar cases, most American authors, especially those 
concerned with biology rather than nomenclature, from 1918 to date 
have followed Walker not because they had critically checked the 
generic names used by him, but simply because it was the most recent 
comprehensive classification outline for North American fresh-water 
shells. As stated elsewhere, this incorrect usage followmg Walker 
dates only from Walker, and has only 33 years standing, which is not 
enough to even raise any questions of usage over priority of names. 

For the record Chenu states (Haldeman, 1847-1848, p. 1, footnote) 
that he and not Haldeman put the name Leptozxis of Rafinesque on 
Haldeman’s pictorial monograph of the genus. Thus Chenu, and 
Pilsbry by formally designating praerosa as genotype in 1917, com- 
pletely cleared up all doubts, concerning Rafinesque’s genus, which had 
existed prior to 1848. 


Genus Eurycaelon Lea, 1864 


Eurycaelon of Lea (1864, p. 3) has as genotype Hurycaelon anthonyr 
(Redfield) by subsequent designation by Walker (1918, p. 36). The 
two known species, Hurycaelon crassa (Haldeman) 1841 and LE. 
anthonyi (Redfield) 1854 are confined to the larger rivers of the upper 
Tennessee drainage as reported by Goodrich (1931). In this con- 
nection must be mentioned the fact that the two names eristata 
Anthony (young specimens) and anthonyi Redfield (adult specimens) 
were published simultaneously in April 1854, according to the printed 
signature dates of volume 6 of the Annals of the Lyceum of Natural 
History of New York. Goodrich, acting as first reviser, selected 
anthony Redfield, on page 130 of volume 6 of the above Annals, over 
the name cristata of Anthony, on page 108, which he placed in syn- 
onymy. In this way Goodrich avoided a confusing change of name 
of the genotype. 


Genus Pleurocera Rafinesque, 1818 


Pleurocera of Rafinesque became monotypic in 1820 with the valid 
publication by Rafinesque of Pleurocera verucosa. Tryon in 1864 and 
1873 did not include the genotype in his usage, so his usage was bio- 
logically and nomenclatorially wrong. Pilsbry in 1896 questioned the 
genotype. Hannibal (1912, p. 169) formally designated the mono- 
type species verrucosa Rafinesque as the genotype, being followed cor- 
rectly by Pilsbry in 1917. 

Bryant Walker, in his 1918 classification, continued to use Tryon’s 
incorrect name for the genus when he used only the second available 


OLD AND NEW WORLD MELANIANS—MORRISON 363 


name, Lithasia Haldeman (1840), which has the species geniculata of 
Haldeman for genotype. 

The smooth species from the Kentucky River named Ellipstoma 
zonalis by Rafinesque in 1818 can only be the species commonly 
known as Lithasia obovata Say 1829. With over 10 years priority, the 
specific name of Rafinesque must be used. Animals of Pleurocera 
zonalis Rafinesque personally collected from the Kentucky River 
have been examined and sketched. The female egg-laying characters 
are illustrated in figure 4 of plate 11. 


Subgenus Ellipstoma Rafinesque, 1818 


In spite of the extreme brevity of Rafinesque’s original description 
(1818b, p. 42), critical reading indicates that this name applies only 
to the group afterward named Angitrema by Haldeman (1841a). 
Hannibal (1912, p. 168) formally designated Rafinesque’s first species, 
Ellipstoma gibbosa, as the genotype, although he misidentified that 
species. When one approaches the identification of gibbosa from 
every angle it is evident that there is only one kind of snail in the 
Ohio and Wabash Rivers with “‘a large knob behind the outward lip.” 
This is the same species named;almost_ three years later as Melania 
armigera Say (1821, p. 178). 

Critical examination of the animal characters of the genotype 
Pleurocera (Ellipstoma) gibbosa Rafinesque is needed to check the bio- 
logical distinction of Eillipstoma as a subgenus, a separate genus, or a 
synonym of Pleurocera, sensu stricto. 


Genus Anaplocamus Dall, 1895 


Anaplocamus of Dall (1895, p. 8), mistakenly described as an Alaskan 
marine shell because of accidentally transposed locality labels, is a 
monotypic genus based on the species ‘“‘Anculosa” dilatata Conrad 
(Rehder, 1942, p. 49). This species of the Kanawha River drainage 
and the upper Cheat River of the Allegheny drainage is the northern 
representative of the Jo group. It has the columellar shell structure 
of Jo, not that of Leptoxis. From a broad biological viewpoint it 
may be considered a subgenus of Jo, or, more conveniently, a separate 
genus closely related thereto. 


Genus Io Lea, 1831 


The genus Jo of Lea is well known; it includes the largest living 
United States Pleuroceridae known. How many species other than 
the nominal genotype fluvialis (Say) there are or were in existence 
must await studies of the animals to check the magnificent analyses 
of shells by Adams (1915). Haldeman, in the American edition of 


364 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Heck’s Iconographic Encyclopaedia (1851, p. 84), has given a general 
description of the animal of Jo: 

Its characters and habits are not those of Fusus, but of [Orytrema] proper, as 
distinguished from Leptozis; for although it inhabits the rapids as well as quiet 
water, in both cases it avoids the current by seeking shelter beneath shelving 
rocks, or in hollows or crevicesin them. The head is large, and with the tentacles 
much exposed; the foot is’as large“as in [Oxytrema]; the* coloration is the same 
(black lines upon an orange ground); the operculum is subspiral as in [Oxytrema], 
the mantle extends into the canal of the shell, but does not form a closed siphon; 
the vent is upon the right side; the mouth is a longitudinal slit; the eyes (which 
are sensitive to the light) are upon a short enlargement of the outer base of the 
tentacles, which may be a little longer than in [Orytrema], and they are not visibly 
annulated. It differs from Fusus, and resembles [Oxytrema] in living upon vege- 
table food; and it moves along in a sluggish manner, moving the head from side 
to side upon the bottom. Jo spinosa and J. tenebrosa are merely varieties of J. 
fluvialis. The spinose individuals are much the most abundant, although the 
species is rare when compared with various species of [Oxytrema]. 


We have taken the liberty of replacing the name “‘Melania”’ in the 
foregoing quotation with that of Oxzytrema, since Haldeman had 
studied and published as typical of ‘‘ Melania’’ the animal of Oxytrema 
virginica (Gmelin) (Haldeman, 1841b, p. 21). 

In this connection, it must be mentioned that the shells of the group 
of ‘‘Pleurocera’”’ alveare Conrad (1834, p. 54, pl. 4, fig. 7) look sus- 
piciously like normal sized representatives west of and downstream 
from the megasomatic Jo, sensu stricto, species. The generic name 
Megara H. and A. Adams (1854, p. 306) has not hitherto been geno- 
typed. Walker (in litt.), quoted by Goodrich (1928, p. 2, footnote 5), 
incorrectly reported that Hannibal (1912, pp. 169, 179) had designated 
the type of Megara. Reference to these pages shows that Hannibal 
listed parts of the original Megara twice, with two different species as 
examples (not as types) in the two synonymy lists 10 pages apart. 
The genotype here designated is the first species, Megara alveare 
(Conrad) 1834. Animal characters of this group remain a desideratum. 
If the group is most closely related to Oxytrema, as it has usually been 
regarded, Megara will remain as a synonym of Oxytrema. If, however, 
these species should prove upon examination to be closest to Jo in 
relationship of animal characters, their group name Megara will be 
available. 


Subfamily PLEUROCERINAE (in Central and South America) 


Genus Pachychilus Lea, 1851 


This genus seems to be the ecological replacement of Oxytrema, to 
the southward of the United States. In my opinion, all the known 
species of this family and subfamily from Central America and the 
West Indies belong to Pachychilus. In this connection, the ichthyol- 


OLD AND NEW WORLD MELANIANS—MORRISON 365 


ogist S. E. Meek should be given credit as the first known discoverer 
of the eggs of any member of the family Pleuroceridae. Reporting on 
collections made in 1906 (Meek, 1908, pp. 205-206), he says: 


Sphaeromelania largillierti Phil. This is the largest and by far the most abundant 
species of shell found in the lake. It is found everywhere along the shore on plants 
and rocks. This species deposits its eggs in January and February. The eggs are 
very large and are usually attached to the under side of rocks. This species is so 
abundant that its large gelatin-like eggs would furnish a considerable amount of 
food for small fishes. It is a very abundant species in Guatemala and San Salvador 
to Central Nicaragua. 


Goodrich and Van der Schalie (1937, pp. 39, 41, 42) have recorded and 
described the eggs of three additional species of Pachychilus as follows: 

Pachychilus glaphyrus (Morelet) 1849. The eggs were found attached to dead 
leaves, which were abundant in the bottom of the arroyo. The leaves were in 
only a few inches of water on a mud-silt bottom near shore. The eggs occur in 
groups of 3 to 5, are round and glassy in appearance, and are irregularly and line- 
arly arranged as a single loosely attached layer. As might be expected in a 
species as large as glaphyrus, the eggs are also large, having a diameter of 3.9 mm. 
(Arroyo Yalchactila of the Rio de la Pasion, about 4 mi. 8. W. of La Ceiba, Alta 
Vera Paz, Guatemala) . 

Pachychilus corvinus (Morelet) 1849. Egg masses of this species were found 
from which the infant shells were emerging. These were smooth, pinkish carnelian, 
translucent, and bluntly rounded at the periphery. The whorls varied from 3 to 
3}. With growth the shell generally becomes black, even though the adults may 
be of a mahogany or yellowish color on the body whorls... . 

Pachychilus pilsbryi Martens 1897. Eggs of this species were found on the lower 
surfaces of stones in a current near shore. The egg masses occur as flat layers 
spread out over flat objects. The size of the egg mass varies, the number of eggs 
in some of them observed running from 8 to 20. The eggs themselves are round, 
shotlike, and transparent. The whole mass is held together by a gelatinous 
coating. The average size of the individual eggs is 2.8 mm. 


We are indeed fortunate in having on record, in the first four species 


so known, the egg-laying characters of three subgenera of the genus 
Pachychilus. 


Subgenus Oxymelania Crosse and Fischer, 1892 


Oxymelania Crosse and Fischer (1892, p. 328) possesses the species 
schiedianus Philippi as genotype by original designation. This group 
of small to medium-sized species has been most completely studied 
in the San Luis Potosi region by Pilsbry, following the collections 
made by A. A. Hinkley. In this subgenus, some species of which are 
widely different in height of spire, and thus of shell outline, the single 
general identifying character is the possession or prominence of macro- 
scopic spiral lirations on the adult shell. The egg-laying habits for 
the species pilsbryi Martens are quoted above. The synonyms of 
Oxymelania include Potamanaz Pilsbry (1893, p. 340), named only a 
few months later, and Lithasiopsis Pilsbry (1910, p. 47). In spite of 

273553542 


366 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


the great similarity of shells, Cubaedomus does not belong here. Ani- 
mal characters prove it to belong in the family Thiaridae, as recorded 
below. Oxymelania is known to range geographically from the Panuco 
River system of México to the headwaters of the Usumasinta River 
in Guatemala. 


Subgenus Pachychilus Lea, sensu stricto, 1851 


Pachychilus Lea (Lea and Lea, 1851, p. 179) originally included 
only two specific names, cumingii Lea and laevissima Sowerby. In 
January 1860, Reeve (1859-1861, pl. 18, fig. 126) designated laevissima 
as the genotype. On the other hand, Reeve’s “laevssima’’ was a 
mixture of the species from Chiapas, México, which has since been 
mistakenly called ‘“Uaevissima’”’ by many authors, and the true laevis- 
sima of Sowerby and Lea from La Guayra, Venezuela. Since the 
mixture of Reeve contained in synonymy the originally included 
laevissima of Sowerby, this designation was valid. Thus the genotype, 
correctly figured on plate 19, figure 133c, of Reeve (1859-1861) is 
the only known South American species of the genus. The typical 
group in Central America is the corvinus or largilliertt group of species, 
characterized by medium- to large-sized shells with a generally elongate- 
ovate or elongate-conic outline, rather flat-sided whorls, and almost 
completely smooth macroscopic sculpture. The egg-laying habits are 
quoted above. The Cuban species nigrata Poey 1858, fuentes Aguayo 
1936, and violaceus Preston 1911 apparently belong here. Synonyms 
of Pachychilus, sensu stricto, include Cercimelania Crosse and Fischer 
(1892, p. 327) with P. liebmanni (Philippi) as genotype, and Sphaero- 
melania Rovereto (1899, p. 109) with the same genotype as Pachy- 
chilus, since it was proposed as a substitute name. Apparently both 
Crosse and Fischer and Hannibal did not know of Reeve’s type desig- 
nation and incorrectly listed P. graphium (Morelet) as the genotype. 


Subgenus Glyptomelania Crosse and Fischer, 1892 


Glyptomelania Crosse and Fischer (1892, p. 328) is confined to 
Central America. This group of large shells with variable to nodose or 
subspinose sculpture includes glaphyrus (Morelet), the subgenotype 
by original designation. The eggs of this species have been recorded 
by Goodrich and Van der Schalie as quoted above. 


Subgenus Pilsbrychilus Morrison, 1952 


Pilsbrychilus Morrison (1952, p. 7) is distinguished from other known 
members of the genus by the markedly sinuous lip-margin developed 
in mature adult shells. It seems probable that examination of the 
animal will serve to corroborate the distinction of this group now based 


OLD AND NEW WORLD MELANIANS—MORRISON 367 


solely on shell characters. The genotype by original designation is 
Pachychilus (Pilsbrychilus) dalli Pilsbry, which was first validly 
published April 17, 1896, in Science (vol. 3, p. 608) ahead of the com- 
plete description by Pilsbry in Proceedings of the Academy of Natural 
Sciences of Philadelphia, June 16, 1896 (p. 269). Fortunately, this 
earliest validation preceded the typographical error ‘‘walli,’’ which 
appeared in Zoologischer Anzeiger in May 1896. 


Genus Doryssa H. and A. Adams, 1854 


Doryssa H. and A. Adams (1854, p. 304), whose genotype is D. 
atra (Bruguiére) from Suriname, ecologically and geographically 
replaces Pachychilus in northern South America from Venezuela to 
the Guianas and Brazil. In both these genera the females have a 
well-developed genital groove ending in the characteristic pleurocerine 
glandular egg-laying sinus near the right edge of the foot. The 
males lack this groove. Doryssa consolidata (Bruguiére) was per- 
sonally collected by the hundreds from rocks in the Mazaruni and 
Cuyuni Rivers of the Essiquibo system, British Guiana, in the 
summer of 1925. At that time the shells were cleaned and the 
radulae (see Baker, 1930, p. 30) extracted from the animals for future 
study of variation. Although at that time I did not know what char- 
acters were critical for subfamily classification, no difference of gross 
anatomy was observed between this British Guiana Doryssa and the 
North American species Oxytrema semicarinata (Say), already crit- 
ically and completely personally studied near Lexington, Ky., in 
1924. The egg-laying season of Doryssa is apparently still unrecorded. 
It was not observed during June, July, or August in the Kartabo 
region. Sheppardiconcha Marshall and Bowles (1932, p. 3), described 
as a new genus from the (Miocene?) fossil deposits of the Upper 
Amazon region of Ecuador, is not a subgenus of Aylacostoma. Refer- 
ence of the group by later authors to Aylacostoma (=Hemisinus) is 
primarily due to the difficulty of complete separation of the genus 
Doryssa of the Pieuroceridae from Aylacostoma of the Thiaridae by 
shell characters alone. According to the shell characters of the 
original material, Sheppardiconcha is a synonym of Doryssa. 


Family PLEUROCERIDAE (in Asia) 
Subfamily PLEUROCERINAE 
Genus Oxytrema Rafinesque, 1819 


In the present state of our knowledge it is no longer possible to 
maintain the generic distinction of certain eastern Asiatic species 
from the genus Oxytrema, which has previously been considered 


368 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


wholly North American. The generic name Hua, proposed by S. F. 
Chen (1943, p. 21) for the group of ‘“Melania”’ telonaria Heude 1888, 
is a synonym of Oxytrema Rafinesque 1819. As is the case in both 
the Appalachian and Rocky Mountain regions of North America, 
the genus Oxytrema is represented in eastern Asia (Korea, China, 
and Thailand) by some species that are generally smooth (Hua) 
and some with strongly nodose or reticulate sculpture. Heude 
figured the female pleurocerine external anatomy of Oxytrema in 
1890 in his excellent work on the Chinese fresh-water mollusks. 
Unfortunately, he misunderstood the structures of the females of 
such species as ‘‘Melania’’ jacquetiana Heude and called them male 
individuals. At present, Heude’s mistake is easy to explain. The 
(female) egg-laying apparatus of these species (Heude 1890, pl. 43, 
fig. 5) is superficially identical in appearance to the (male) intromittent 
organ of certain operculate land snails of the subfamily Cyclophorinae 
(family Cyclophoridae) (see Heude 1890, pl. 42, fig. 12c), as a com- 
parison of Heude’s own figures will demonstrate. It is this extreme 
similarity in appearance that led Heude to an understandable mis- 
interpretation and has effectively hidden the complete understanding 
of his critical discoveries for many years. 

The present studies have corroborated those of Heude. The 
animals of Oxytrema telonaria (Heude) 1888 are dioecious; the sexes 
may be distinguished by the presence or absence of the female genital 
groove on the right side of the foot. There is a glandular, egg-laying 
pit at the terminal end of the groove a little above the right edge of 
the foot. The epithelium lining this groove is markedly different 
from that of the adjacent area and is easily distinguished in preserved 
material. The posterior edge of the groove is projected into the pit 
as a sort of papilla at its termination. The oviduct is not enlarged 
into a uterus; there are no embryos in the body behind the head. 
Animals of the following Asiatic species have been personally examined 
and found to exhibit the same pleurocerine reproductive anatomy 
with but little difference in minor details: 0. nodifila (Martens) 1886, 
Han River, Korea (see pl. 11, fig. 3); O. towcheana (Heude) 1888, 
Min River, Fukien, China; 0. jacquetiana (Heude) 1890, Shaochsing, 
Chekiang, China; O. peregrinorum (Heude) 1890, Fukien, China; 
O. moutoniana (Heude) 1890, Sa-Hsien, Fukien, China; O. joretiana 
(Heude) 1890, Chekiang, China; and O. bazlleti (Bavay and Dautzen- 
berg) 1910, Szechwan, China. How many of the other Asiatic species 
listed by S. F. Chen in these groups will prove to belong to Orytrema 
can be determined only when their reproductive anatomy is examined. 
The presence of Oxytrema species in the recent fauna on both sides of 
the North Pacific Ocean demonstrates a geographic distribution similar 
to that of the genera Viviparus and Anodonta, sensu stricto. It is 


OLD AND NEW WORLD MELANIANS—MORRISON 369 


evident also that the western American (Alaskan) region consti- 
tuted the pathway of migration of these fresh-water mollusks between 
Asia and North America. 


Genus Paludomus Swainson, 1840 


With a short, ovate shell-shape corresponding to that of the Ameri- 
can genus Leptozis, the Asiatic genus Paludomus (Swainson, 1840, 
pp. 198, 340) also belongs to the subfamily Pleurocerinae. The 
anatomy of the animals of Paludomus tanschaurica (Gmelin) has 
been completely studied by Seshaiya (1934). The finding by Seshaiya 
of a tubular structure which he called a “penis” along the terminal 
portion of the primary gonoduct of the male must be discounted as 
a glandular discovery until it is proven to be an intromittent organ. 
As Seshaiya himself pointed out, it is not muscular as is the case 
of the similarly located and functional male organ of the Tiphobiimae. 
Seshaiya believed these animals oviparous, as he found no evidence of 
Ovoviviparity in their anatomy. He did not, however, mention either 
the egg-laying groove or the pit. Paludomus labiosus (Benson) (?) 
from Thailand and Paludomus maculatus Lea from India (also 
members of the subgenus Paludomus, sensu stricto, with its concentric 
operculum) have been personally examined. The female animals are 
pleurocerine, with an egg-laying groove and pit in the side of the foot 
and without any brood pouch (see pl. 11, fig. 6). The true biological 
relations of the other subgenera such as Hemimitra, which differ 
markedly in shell and opercular characters, are still unknown or 
unrecorded. 


Subfamily LAVIGERIINAE Thiele, 1929 


Members of the subfamily Lavigeriinae Thiele (1929, p. 79) show 
the full dioecious mode of reproduction of the family. That is, there 
is a normal 1:1 sex ratio of males to females as in all other known 
members of the Pleuroceridae. The most important distinguishing 
character is the possession of a uterine brood pouch similar to that in 
the Viviparidae. As in the typical subfamily (Pleurocerinae), the 
males possess no intromittent structures whatsoever. Coincident to 
the development of ovoviviparity, the females have lost (at least do 
not possess) the egg-laying sinus and papilla characteristic of the 
Pleurocerinae. The name Semisulcospirinae Morrison (1952, p. 8) is 
a synonym of the earlier subfamily name Lavigeriinae. 


Genus Semisulcospira Boettger, 1886 


The genotype of Semisulcospira Boettger (1886, p. 4) is the well- 
known Japanese species Semisulcospira libertina (Gould) 1862... Ex- 
amination of many specimens from the Hayakawa;River, Honshu, 

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370 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Japan, sent to the U.S. National Museum a few years ago by Captain 
Honess, U. S. Army Sanitary Corps, conclusively demonstrated that 
this species is Ovoviviparous, as previously suspected. The head and 
snout of these snails are uncomplicated in either sex by external re- 
productive structures of any kind. Coitus is probably accomplished 
by apposition of the mantle edges, with perhaps some prolapsis of the 
genital duct openings. The lower part of the female reproductive 
duct (oviduct) is enlarged to form a functional uterus. It is thin- 
walled when fully distended with young and apparently is not glandu- 
lar. The hundreds of embryonic young of uniformly small size 
developed at one time appear to be packed in axial rows, visible ex- 
ternally when the animal is removed from the shell and resembling 
the conglutinates of glochidia of the North American fresh-water 
mussel genus Strophitus. In some cases the young nearest the ovi- 
duct are not fully developed, showing that development of the ova is 
at least partly progressive as the young approach the uterine mouth. 
On the other hand, the great uniformity of size of all the fully de- 
veloped young, with shells of about 2 whorls, would indicate a definite 
seasonal, mass release of the young. At any rate, there is no indica- 
tion of any continued growth to large size of a few embryos at a time 
or continued parturition over a long season or throughout the year 
as in certain genera of the family Thiaridae. 

Recently, Corp. W. E. Old of the U. S. Army Signal Corps sent 
four species of ‘“‘Melania’” from Korea to the U. S. National Museum. 
Examination of the animals by both R. T. Abbott and the author has 
proven that “Melania” nodifila Martens is an Oxytrema as noted above, 
while the species gottschei Martens, nodiperda Martens, and graniperda 
Martens (1905) are members of the genus Semisulcospira. The 
geographic range of Semisulcospira across continental Asia is imper- 
fectly known and in need of continued research. 


Family PLEUROCERIDAE (in Africa) 
Subfamily PLEUROCERINAE 
Genus Potadoma Swainson, 1840 


The genotype of Potadoma Swainson (1840, pp. 200, 341), P. freethiz 
(Gray), must be examined to completely clear up the position of this 
genus. Pilsbry and Bequaert (1927) have reported that these snails 
are apparently oviparous, with the edge of the mantle smooth and even. 
They compare these species to some species of the American genus 
Pachychilus. If Potadoma is oviparous as reported, with the egg-laying 
groove, it belongs in the subfamily Pleurocerinae, and the subfamilies 


OLD AND NEW WORLD MELANIANS—MORRISON 371 


Potadominae Pilsbry and Bequaert and the Paludominae are included 
synonyms. In addition to the species called Potadoma by Pilsbry 
and Bequaert, some of the species they called ‘“‘Afelanoides”’ also belong 
to Potadoma of Swainson. ‘‘Melanoides’’ species such as ignobilis 
(Thiele) (Pilsbry and Bequaert, 1927, p. 278, pl. 26, figs. 1, 2) are 
smooth; others such as medjeorum Pilsbry and Bequaert (1927, p. 283, 
fig. 47) and tornata (Martens) (Pilsbry and Bequaert, 1927, p. 283, fig. 
48) possess strong spiral sculpture. Still others, such as wagenia Pilsbry 
and Bequaert (1927, p. 262, pl. 22, figs. 14-16), crawshayi (EK. A. Smith) 
and mweruensis (EK. A. Smith) (Pilsbry and Bequaert, 1927, pp. 264— 
265, pl. 22, figs. 19-22) show strong plicate to cancellate shell sculpture. 
The total picture of sculpture in this genus seems exactly parallel to 
that seen in Appalachia, western America, and eastern Asia in the 
genus Oxytrema. 


Genus Limnotrochus EK. A. Smith, 1880 


1880. Limnotrochus E. A. Smith, 1880b, p. 425. (Genotype, Limnotrochus 
thomsont, E. A. Smith, 1880.) 


Genus Paramelania E. A. Smith, 1881 


1881. Paramelania KE. A. Smith, 1881b, p. 559. (Genotype, Paramelania 
damont E. A. Smith, 1881, by subsequent designation by Pilsbry and 
Bequaert, 1927, p. 320.) 


Genus Spekia Bourguignat, 1879 


1879. Spekia Bourguignat, 1879, p.27. (Genotype, Spekia zonatus (Woodward), 
1859, by monotypy.) 

Pilsbry and Bequaert have said that, as far as known, the three 
above-named genera are oviparous. If complete study of the animals 
confirms this and reveals the characteristic egg-laying pit in the side 
of the foot, these Lake Tanganyika forms are also Pleurocerinae. 
Moore (1899a, p. 171) found neither an external groove nor a brood 
pouch of any type in the females of Spekia zonata (Woodward) 
examined by him. This condition seems identical in appearance to 
that of those North American members of the pleurocerine genus 
Oxytrema, in which the female egg-laying groove is not demarcated 
structurally and can only be seen functionally as a linear depression 
in the skin during the breeding season. 


Subfamily LAVIGERIINAE Thiele, 1929 


The statement of Pilsbry and Bequaert (1927, p. 300) in regard to 
the ovoviviparity of the mixture of species they called ‘‘ Melanoides’’ is 
in need of clarification. They left the problem unsolved when they 
said: “Typhobia, Bathanalia, and Lavigeria are viviparous, having the 


372 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


last part of the oviduct enlarged to form a brood pouch in the mantle, 
as in Melanoides and Viviparus.’”’ The fact is that Melanoides does 
not have a uterine brood pouch as in the family Viviparidae. ‘True 
Melanoides species have the subhaemocoelic brood pouch in the back 
of the neck, with the brood pouch pore on the right, as in all other 
members of the Thiaridae. 

If Pilsbry and Bequaert actually saw uterine brood pouches in the 
animals of any African species of ‘‘Melanoides,’”’ they probably were 
dealing with members of the subfamily Lavigeriinae, of an unnamed 
genus. In this connection, the possibility should be stated frankly 
that the subfamilies Lavigeriinae and Tiphobiinae may in the future 
prove to be biologically confluent. Much more study of the animals of 
African fresh-water species both in and out of Lake Tanganyika is 
needed to prove or disprove this possibility. As defined above, the 
subfamily Lavigeriinae includes those genera of pleurocerid snails in 
which the males have no intromittent organ and the females possess 
uterine brood pouches. 


Genus Lavigeria Bourguignat, 1888 


The genus Lavigeria Bourguignat (1888, p. 33) is another of the 
peculiar pleurocerid forms found in Lake Tanganyika. In addition to 
the shell characters, this genus differs from others by its operculum, 
which is modified by partial uncoiling to a subspiral form. Moore 
1899b, p. 192, pl. 20, fig. 6) has furnished anatomical studies of the 
species Lavigeria coronata Bourguignat. He called the animal ‘‘Nas- 
sopsis nassa Woodward,” while Pilsbry and Bequaert have referred 
this anatomical study to the genotype which they designated, namely: 
Lavigeria grandis (EK. A. Smith) 1881. Moore’s figure of the shell 
proves he studied the species coronata. He found that this genus 
possesses a uterine brood pouch similar to that of Viviparus and of 
the Tiphobiinae. According to him, the males do not possess any 
secondarily developed intromittent organ such as is found in the 
mantle edge of the males of Tiphobia. 


Genus Bourguignatia Giraud, 1885 
1885. Bourguignatia Giraud, 1885, p. 193, pl. 7, figs. 5-7. (Genotype, Bourguigna- 
tia imperialis Giraud, 1885, by monotypy.) 
Genus Edgaria Bourguignat, 1888 


1888. Edgaria Bourguignat, 1888, p. 38. (Genotype, Edgaria paucicostata (HE. A. 
Smith), 1881, by subsequent designation by Pilsbry and Bequaert, 1927, 
p. 328.) 


The shells of the Bourguignatia, Lavigeria, Edgaria, and Paramelania 
complex of Lake Tanganyika are almost identical in general characters; 
however, there are two distinct types of opercula known in the group. 


OLD AND NEW WORLD MELANIANS—MORRISON 373 


Our present lack of anatomical knowledge of all but Lavigeria makes 
it impossible to say whether two or more subfamilies are represented 
in this complex of shells inhabiting the rocky shorelines of Lake 
Tanganyika. 

Genus Bythoceras Moore, 1898 


1898. Bythoceras Moore, 1898c, p.452. (Genotype, Bythoceras iridescens Moore, 
1898, by monotypy.) 

These shells from deeper waters of Lake Tanganyika are also very 
similar to those of the Lavigeria complex; their opercula differ from 
those of Lavigeria in being secondarily concentric around a paucispiral 
nucleus. The general statement by Moore (1898d) that Bythoceras 
is much more like Tangenyicia in anatomy than (‘‘ Nassopsis’=) 
Lavigeria is in need of corroboration or clarification. Neither 
is it clear from Moore’s subsequent statements and figures (1899b) 
whether Bythoceras is ovoviviparous or not. 

The final allocation of these names and also of Paramelania and the 
subfamily name Paramelaniinae must await the complete recording 
of shell, opercular, and reproductive characters of all members of the 
Lavigeria complex of Lake Tanganyika. If the reproductive anatomy 
proves identical, Paramelaniinae (Moore, 1898b, p. 315) will super- 
sede the name Lavigeriinae (Thiele, 1929, pp. 79, 83) used at this time. 


Subfamily TreHoBIINAE Moore, 1898 


These are pleurocerid snails modified in the males by the develop- 
ment of a secondary, eversible ‘‘penis’” or vergic structure in the 
mantle edge near the end of the vas deferens. As far as known, these 
are the only ‘“Melanian” snails of any kind that possess any intro- 
mittent structures whatsoever. The females are ovoviviparous, with 
the terminal portion of the oviduct modified into a uterine brood 
pouch for the young. At present, our knowledge—or lack of 
knowledge—indicates that the Tiphobiinae are solely African in 
geographic distribution, parallel to but not identical with the Asiatic 
and African subfamily Lavigeriinae. 


Genus Tiphobia E. A. Smith, 1880 


The genotype of 7iphobia HE. A. Smith (1880a, p. 348, pl. 31, fig. 6), 
T. horei E. A. Smith, 1880, possesses the reproductive characters of 
the subfamily, as defined above, according to the studies of Moore 
(1898a). The statement of Pilsbry and Bequaert that the characters 
of the Tiphobiidae do not distinguish them from the Melaniidae is 
based on their misunderstanding of the true nature of the brood 
pouch of the Thiaridae (‘‘Melaniidae”’). Tiphobia is one of the 
endemic genera of Lake Tanganyika, with a spinose shell and a 


374 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


secondarily concentric operculum. Its complete biological relation- 
ship with the other Tanganyika genera and with extra-Tanganyika 
forms must be clarified by future studies. 


Genus Bathanalia Moore, 1898 


The genotype, Bathanalia howesi Moore, 1898, is stated by Moore 
(1898a) to be “almost identical anatomically with Tiphobia horet.” 
This genus differs from Tiphobia by shell characters such as height of 
spire and the perforate columella. In addition, Bathanalia possesses 
an unmodified (paucispiral) operculum. 


Family THIARIDAE 


The pattern of reproduction is the same for all known members of 
the family Thiaridae. A high degree of specialization in this one 
character is indicated by this observed pattern of ovoviviparous 
parthenogenicity. In other words, there is no male individual known 
in this entire family of snails. To my knowledge, the parthenogenesis 
of the Thiaridae has not been examined cytologically to determine 
whether it is haploid or diploid. 

There is required only one individual (any individual) to start 
a new population or colony if it reaches a new fresh-water locality by 
stream capture or by adventitious transportation. The resultant 
ability of these snails theoretically to spread more rapidly may partly 
explain their wider distribution in suitable habitats on islands in both 
the Pacific and the West Indies and their wider expansion southward 
across South America than that exhibited at present by the family 
Pleuroceridae. Certain corollaries are assumed in this theoretical 
explanation of the differences and peculiarities of distribution of the 
two fresh-water families of Pleuroceridae and Thiaridae. Structurally, 
the animals of these families are relatively primitive, exceedingly so 
as regards their fundamental reproductive characters. Without any 
proof to the contrary, we may assume them to be of approximately 
equal antiquity, geologically speaking. Reasoning from the special- 
ized pattern of reproduction of the Thiaridae, we may logically argue 
that they are the younger group evolved from the older ancestral type 
represented now in fresh water by the Pleuroceridae. If this be true, 
the younger has outstripped the more primitive older type in geo- 
graphic expansion over geologic time from the era of their differentia- 
tion up to the present. If the Thiaridae were ever in North America 
we must assume that they are no longer represented there because the 
conditions of existence are not now or were not, during some previous 
geologic era, favorable to their continued existence in those northern 
areas. Their expansion across the North American continent may be 


OLD AND NEW WORLD MELANIANS—MORRISON 375 


unrepresented in any fossil record, known or yet undiscovered, be- 
cause successfully living fresh-water mollusk species seldom undergo 
fossilization. They usually enter the fossil record only when a 
habitat change brings about the extinction of populations or species. 
On the other hand, the present geographic distribution of the Thiaridae 
argues for their greater ability to spread across oceanic areas on island 
stepping stones without the necessity for any continuous “land 
bridges.” 

There is one important taxonomic problem present in the Thiaridae 
that has not often been acknowledged in the study of these or any 
other parthenogenetic animal species. We know by observation the 
great variability of individuals within the species of the dioecious 
type such as the Pleuroceridae. This variability is the greatest 
single factor contributing to confusion in past and present studies of 
these fresh-water shells. The variability of individuals of the related 
Thiaridae may be either increased or decreased because of their 
reduction to unilateral ancestry. What we actually have in the 
entire family of the Thiaridae is the possibility of clone formation in 
every existing population of these animals. These is no biparental 
or cross-inheritance control or check on individual variation in their 
present parthenogenetic state. Wise indeed is the scientist who can 
tell whether a clone is a species or not, and be right every time, in the 
case of the Thiaridae. 


Family TutaripaE (in the Americas) 


The earliest generic name for any of the indigenous American 
species of this family is Aylacostoma Spix (1827, p. 15, pl. 8). This 
manuscript name of Spix was first published in the synonymy of both 
Melania tuberculata Wagner (= Aylacostoma tuberculatum Spix) and 
Melania scalaris Wagner (= Aylacostoma glabrum Spix) in connection 
with the species description, and also on plate 8 (Spix, 1827) without 
the indication of synonymy. ‘This is in the first edition (Munich) of 
plates, not the second edition (Leipsic). As a manuscript name of 
Spix, published in direct connection with two validly described and 
figured species, Aylacostoma is valid and available. The genotype, 
by subsequent designation by Morrison (1952, p. 8), is Melania 
scalaris Wagner (= Aylacostoma glabrum Spix). This species, as 
comparison of the figures will demonstrate, is that form called behnii 
by Reeve (1859-1861, pl. 2, fig. 8) in his 1860 monograph on Hemisinus. 

Swainson (1840, pp. 200, 341) was next to furnish a name for Ameri- 
can Thiaridae. Hemisinus is his generic name given to the species 
lineolata Gray from Jamaica. The later emendations, Aulacostoma 
Agassiz 1846 and Semisinus Crosse and Fischer 1885, are both 


376 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


unnecessary and invalid under present international rules of nomen- 
clature. Basistoma Lea (1852, p. 295) is exactly equivalent biologi- 
cally to Aylacostoma, being based on the same type of shell from 
South America. Verena H. and A. Adams (1854, p. 308) differs from 
the typical Aylacostoma group in shorter shell outline and in sculpture. 


Genus Cubaedomus Thiele, 1928 


This genus (Thiele, 1928, p. 401), based on the species brevis Orbigny 
from Cuba, is the only American group clearly and generically distinct 
from Aylacostoma on shell characters. The shell is ovate-conic, with 
a trace of nodulous coronate sculpture on the upper slope or shoulder 
of the body whorl. There is no columellar notch or sinus in the 
aperture, which is evenly rounded below (anteriorly). The operculum 
is paucispiral. Animals of this species, dried in the shells, were 
recently boiled up in water (softened) and personally examined. 
These specimens (USNM 407991) were collected by Jaume from Rio 
“Los Cayos”, Bahia Hondo, Pinar del Rio, Cuba. The mantle is 
fringed. ‘The embryonic young are relatively large, and there were 
only two in the subhaemocoelic brood pouch in the neck region at one 
time in the material examined (pl. 11, fig. 19). Proof that this 
genus belongs to the family Thiaridae gives us another excellent 
example of concurrent and convergent evolution. Both Pilsbry (1893) 
and Aguayo (1944, p. 69) have previously considered the species 
brevis Orbigny of Cuba congeneric with (““Potamanaz’”’=) Pachychilus 
(Oxymelania) pilsbryx (Martens) of the Guatemala region, a member 
of the Pleuroceridae, because of the almost identical shell character- 
istics of the two species. 


Genus Aylacostoma Spix, 1827 


Subgenus Hemisinus Swainson, 1840 


Genotype: (Melania lineolata Gray =) Aylacostoma (Hemisinus) 
lineolatum (Gray) 1828. 

This group of elongate-ovate or ovate-conic shells, almost completely 
lacking macroscopic spiral sculpture, includes one or two species from 
Cuba, the genotype from Jamaica, and a few other species of Central 
and South America from Panamé to Ecuador and Pert. 


Subgenus Aylacostoma Spix, 1827, sensu stricto 


Genotype: (Aylacostoma glabrum Spiz=) Aylacostoma (Aylacostoma) 
scalare (Wagner) 1827. 

This typical group includes one species from the Atlantic side of 
Central America, one species from the Pacific drainage in the Darién 
region of Panama, and numerous species from the Magdalena, 
Orinoco, Amazon, and southern Brazil regions of South America. 


OLD AND NEW WORLD MELANIANS—MORRISON 377 


These shells have spiral sculpture more or less prominent on most of 
of the whorls, and in some of the species there is a peculiar swelling 
or expansion of the body whorl of adults. The female reproductive 
anatomy and embryos (from the brood pouch) of the Central American 
species Aylacostoma ruginosum (Morelet) were figured by Crosse and 
Fischer (1892, pl. 49, fig. 10). Im these figures the brood pouch 
relation is not clearly \indicated; it was not fully understood to be 
a separate adjunct to the primary female reproductive system. 
Hinkley (1920, p. 47) also has recorded the reproduction and ecology 
of this species. He says: 

Their trail was made by burrowing instead of crawling on the surface as with 
other forms. They burrowed somewhat like a mole, and often the little mole-like 
ridge could be followed quite a distance, and the mollusk found working under 
cover. 

It is a viviparous genus. When cleaning these shells the embryos run from one 
to three to the individual. None were noticed with more than three. 
Hinkley’s observations on the embryos are corroborated by dried 
animals, received with the shells (USNM 218018) from Hinkley, 
which show two large young in situ in the brood pouch. These 
specimens are from Lake Ysabal, Jocolo, Guatemala. 


Subgenus Longiverena Pilsbry and Olsson, 1935 


Genotype: (Aylacostoma tuberculatum Spix, 1827=Melania tubercu- 
lata Wagner, 1827, not Melania tuberculata (Miller) 1776=) Ayla- 
costoma (Longiverena) tuberculatum Spix, 1827, by subsequent 
designation by Morrison, 1952. 

This subgeneric group of Aylacostoma species possesses prominent 
longitudinal sculpture in addition to the spiral lirae present on the 
shells of Aylocostoma, sensu stricto. In this way, the upper spire 
whorls may greatly resemble the corresponding portion of the shell 
of some species of Doryssa described from the same general region. 
More study is required for proof, but this general resemblance of 
Doryssa and Longiverena shells may be another outstanding example 
of concurrent and convergent evolution of the families Pleuroceridae 
and Thiaridae. 

Pilsbry and Olsson (1935, p. 11) did not formally designate any of 
the originally included species, recent or fossil, as genotype, so we 
designated the living species tuberculatum in order that the animal 
characters of the group may be used to correctly place this unit in 
the total biological picture. 


Subgenus Verena H. and A. Adams, 1854 


- Genotype: (Melania crenocarina Moricand=) Aylacostoma (Verena) 
crenocarina (Moricand). 


378 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


This fourth South American group, Verena, has been considered 
distinct by most authors. Its principal known distinguishing char- 
acter is the possession of strong, undulate or scalloped spiral ridges 
on the post-nuclear whorls. Dr. Olsson of the Academy of Natural 
Sciences of Philadelphia has recently found species in the fossil record 
from the upper Amazon region that prove the existence of Verena in 
that area for considerable time. These fossil species possess shells 
with sculpture identical in type to that of crenocarina (Moricand), 
but, unlike that living species, show shell outlines more or less com- 
pletely transitional to the high-spired outline of Aylacostoma, sensu 
stricto. 

Genus Tarebia H. and A. Adams, 1854 


One Asiatic species, probably Tarebia lateritia (Lea) from the 
Hawaiian Islands, has recently been introduced to American waters 
with aquarium plants. This species was first discovered living feral 
in Lithia Sulphur Springs, near Tampa, Fla., in December 1947 by 
C. W. Cooke of the U. S. Geological Survey. It was not there in 
March 1940 on the occasion of an earlier complete collection of mol- 
lusk species by Dr. Cooke from that spring. This kind of introduction 
is highly dangerous to public health because this Asiatic type of snail 
is one of the primary intermediate host species of the human lung 
fluke (Paragonimiasis). 


Family THrarmDaz (in Asia) 
Genus Thiara Réding, 1798 


This genus (Réding, 1798, p. 109) was not originally monotypic. 
It included two valid specific names, amarula (Linnaeus) 1758 (Chem- 
nitz, 1786, pl. 134, figs. 1218, 1219) and cancellata Roding 1798 
(Chemnitz, 1786, pl. 134, figs. 1220, 1221), as well as three nomina 
nuda. Hermannsen (1849, p. 576) did not explicitly state that 
amarula Linnaeus was the type. In Latin, he said it was a genus ‘‘of 
the type of.”” However, Brot (1874, p. 7) formally designated amarula 
the genotype. In the same monographic work he named two other 
groups which are synonyms of Thiara. Tiaropsis Brot (1874, p. 7), 
based on the genotype species winterd von dem Busch (1842, p. 1, 
pl. 1, figs. 1,2), and Plotiopsis Brot (1874, p. 7), based on the genotype 
species balonnensis Conrad (1850, p. 11), both possess the same shell 
characters as Thiara, differing principally in height of spire and size 
of shell. At present these differences are only considered of specific 
value. Another recently created synonym is Pseudoplotia Forcart 
1950, with the species scabra Miiller 1776 as originally designated 


genotype. 


OLD AND NEW WORLD MELANIANS—MORRISON 379 


Subgenus Thiara Roding, 1798, sensu stricto. 


Genotype: Thiara (Thiara) amarula (Linnaeus), 1758. 

The animals of Thiara spp. from Szechwan, China, Mindanao, 
Philippine Islands, and Okinawa, Ryukyu Islands, have been exam- 
ined by me and found to possess the reproductive structures described 
above as distinguishing characteristics of the family (pl. 11, figs. 9, 11). 


Subgenus Setaeara Morrison, 1952 


Genotype: (Thiara cancellata Réding 1798=) Thiara (Setaeara) 
cancellata Roding 1798. 

The genotype has been known for many years under the name of 
setosa Swainson because the far earlier name of Réding has not 
hitherto been critically examined. These shells possess stronger 
spiral sculpture than those of Thzara, sensu stricto, and the ‘‘crown” 
of spines on the shoulder of the whorls is continued upward as a set 
of cuticular setae, easily broken off, and almost always missing from 
imperfect shells. This group of only a few species was proposed as 
a subgenus (Morrison, 1952, p. 8) until animal characters are found 
that confirm its separation from Thiara, sensu stricto, by means of 
the shell. 


Genus Tarebia H. and A. Adams, 1854 


Genotype: (Melania semigranosa von dem Busch =) Tarebia semi- 
granosa (von dem Busch) 1842, by subsequent designation by Morri- 
son, 1952. 

Reexamination of the type designation of this genus by Brot (1874, 
p. 7) shows it to be completely inoperative. His statement that 
“‘granifera”’ is the type refers to Melania granifera von dem Busch, 
the only such specific name originally included by H. and A. Adams. 
To the best of my knowledge, 7. granifera H. and A. Adams is still a 
nomen nudum. This nomen nudum is proven not to be the same 
as Melania granifera of Lamarck by H. and A. Adams, who listed 
Lamarck’s specific name separately and distinctly as a member of 
the genus Plotia. One of the valid, originally included species was 
designated as genotype (Morrison, 1952, p. 8) in order to correctly 
and legally fix Tarebia upon the group of thiarid snails to which it 
has been restricted and applied since the time of Brot (1874). 

The animals of JZ. lateritia (Lea) from China, of TJ. granifera 
(Lamarck) from the Philippines, and, more recently, of the Tarebia 
colony accidentally introduced to Lithia Sulphur Springs, near 
Tampa, Fla., have been examined for the reproductive characters 
and proven thiarid. 


380 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Genus Sermyla H. and A. Adams, 1854 


Genotype: (Melania mitra Dunker =) Sermyla tornatella (Lea) 1850. 

Recent examination of the animals of S. kowloonensis S. F. Chen 
1943 and of S. mauwiensis (Lea), from China and the Hawaiian Islands 
respectively, proves this genus typically thiarid, as most authors have 
considered it. These shells possess arcuate, longitudinal sculpture on 
the upper (posterior) portion of the whorls, strongly contrasting with 
the spiral lirae below (anterior). They are relatively short-spired. 


Genus Stenomelania Fischer, 1885 


Genotype: (Melania aspirans Hinds=) Stenomelania aspirans 
(Hinds). 

Stenomelama Fischer (1885, p. 701) is a thiarid genus of elongate 
shells with the spire sometimes exceedingly attenuate and sharp- 
pointed, when not eroded away. Members of this genus possess the 
brood pouch typical of the family, that holds the eggs until they 
develop into veligers. Seshaiya (1940, p. 331) has shown that these 
snails, living in estuarine habitats, release the embryonic young into 
the water in the veliger stage. There is thus a free-swimming larval 
stage in the life history. These observations recently have been 
corroborated in the case of certain Philippine species of the genus by 
R. T. Abbott. The species M. aspirans is the genotype by monotypy. 
Another species, M. hastula (Lea), is the genotype by original desig- 
nation of the synonymous generic name Radina of Preston (1915, 
Dp. LO); 
bey Genus Melanoides Olivier, 1804 


Genotype: (Melanoides fasciolata Olivier=) Melanoides tuberculata 
(Miller) 1776. 

The shells of Méelanoides are elongate-conic, of a number of regularly 
increasing whorls. The spire is usually twice the length of the 
aperture, or more. The sculpture is smooth to finely or coarsely 
tuberculate, in different species, without any material difference 
between the younger and older portions of the shell. The operculum 
is clearly paucispiral, with the nucleus so near the base that in some 
cases it may approach the subspiral condition. 

A personal check on available material in the U. S. National 
Museum collections has shown that the following species belong to 
Melanoides, agreeing exactly in the thiarid type of ovoviviparous, 
parthenogenetic reproduction: M. chinensis (Nevill), China; M. 
surifuensis S. F. Chen, Szechwan, China; 1. turriculus (Lea), Leyte, 
Philippine Islands; Melanoides sp., Rota, Marianas Islands; M. vainafa 
(Gould), Ofu, Samoa; and M. indefinata (Lea), Oahu, Hawaiian 
Islands. 


OLD AND NEW WORLD MELANIANS—MORRISON 381 


Genus Sulcospira Troschel, 1857 


Genotype: (Melania sulcospira Mousson=) Sulcospira sulcospira 
(Mousson). 

Recorded as “‘also viviparous,” this group must be studied further 
before its true relationships to the other genera from the Malayan 
region can be proved. ‘Troschel (1857, p. 114) has recorded the 
radular characters; the shell appears similar to certain species of 
Brotia, to Tylomelania, or to immature individuals of Balanocochlis. 
Until the animal characters are reexamined, Sulcospira may be left 
tentatively in the ““Melanoides complex.” 


Genus Balanocochlis Fischer, 1885 


Genotype: (Melania glans von dem Busch=) Balanocochlis glans 
(von dem Busch) 1842. 

Philippine animals of this species, collected by R. T. Abbott at San 
Ramon, Mindanao (USNM 543951), have recently been examined. 
These animals prove the genus to be thiarid. There is a minute 
brood-pouch pore bordered with ‘‘flaps,” similar to that figured for 
Melanoides turriculus (pl. 11, fig. 14), low on the right side and close 
to the fringed mantle edge. Apparently there are many hundreds of 
minute eggs in the brood pouch. Either they develop as numerous, 
small, equal-sized young shells as in certain Melanoides species, or 
they may be released in the veliger stage for a free-swimming larval 
period, as is known for the genus Stenomelania. The material ex- 
amined, which contained only the eggs, is not conclusive on this point. 
In any case, the shell and opercular characters are distinct enough 
from any other thiarids known to maintain the full generic separation 
of Balanocochlis. The earlier name Melania inermis Lesson 1830 was 
preoccupied by Gray in 1825; the later name M. siccata von dem 
Busch 1843 probably represents young of the species Balanocochlis 

lans. 
: Genus Tylomelania P. and F. Sarasin, 1898 


Genotype: Tylomelania neritiformis P. and F. Sarasin. 

This subpaludomiform group of shells only appears as a separate 
development of shell outline and proportions; they have the same type 
of “‘paleomelanian” operculum and the same general radular char- 
acters as do Antimelania and Brotia. The animals of all the groups 
must be completely and carefully reexamined before the genera (and 
the distinctions of the genera) of the “‘Melanoides complex” of the 
Thiaridae can be properly organized or understood. Shell, radular, 
and opercular characters of this group were carefully recorded by the 
Sarasins (1898, p. 52, pl. 4, figs. 56, 57; pl. 8, figs. 110, 111; pl. 9, fig. 
115), but they did not examine the details of the reproduction of 
Tylomelania. 


382 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Genus Antimelania Crosse and Fischer, 1892 


Genotype: (Melania variabilis Benson 1836 (not Melania variabilis 
Defrance 1823)=) Antimelania costula (Rafinesque) 1833. 

As the well-known but preoccupied and preceded name for the 
genotype implies, the shell sculpture of members of this genus is 
variable. These shells are similar to Melanoides but usually larger in 
size, with the sculpture bolder when present. The operculum is 
paleomelanian, often with about 6 turns in contrast to that of 
Melanoides with about 2 turns. The apex or nuclear whorl of these 
shells is perfectly symmetrical in uneroded individuals. The aperture 
is more-or-less regularly rounded at the base, without a forward 
projecting rounded angle as in Brotia. 

Specimens of Antimelania soriniana (Heude) collected in 1940 by 
Dr. H. T. Chen at Tai-Mo-Shan, Kwangtung, China, possess a paleo- 
melanian operculum (with large nucleus) as in Heude’s figures (1890, 
pl. 41, figs. 6, 6a). They were all ovoviviparous, parthenogenetic 
females containing hundreds of minute young in the brood pouch, 
which is enormously expanded into the body cavity behind the head. 
They possessed a narrow genital groove terminating abruptly on the 
right side of the foot, without any trace of the pleurocerine type of 
ege-laying sinus. 

Apparently Pilsbry and Bequaert (1927, p. 300) did not know that 
the specific name variabilis was preoccupied when they designated it 
the genotype of Antimelania (Crosse and Fischer, 1892, p. 313). A 
search for the correct name to use in replacement led to the discovery 
that costula Rafinesque (1833, p. 166) is clearly recognizable as identi- 
cal to Benson’s variabilis. 

Moore (1899a, p. 161) has recorded the brood-pouch ovoviviparity 
of “Melania” episcopalis (Lea). His figure (pl. 14, fig. 13) of the 
female Antimelania episcopalis (Lea) agrees with our observations on 
the genus Antimelania. The figure given by him of a “male” animal 
of this same species is either a nongravid or nonbreeding female of the 
same species, or a female of the pleurocerine genus Oxytrema with only 
the genital groove and egg-laying pit in the side of the foot. Moore 
(1899a, p. 163) mentions the presence of males in Tanganyicia and 
“‘Melania,”’ but nowhere proves it. The presence of any males in 
this family is still not proved. 


Genus Brotia H. Adams, 1866 


Genotype: (Melania pagodula Gould 1847 (not Melania pagodulus 
Reeve 1860)=) Brotia pagodula (Gould) 1847. 

Shells of this genus are elongate-conic, of variable sculpture, and 
with the columellar angle of the basal lip of the aperture produced 


OLD AND NEW WORLD MELANIANS—MORRISON 883 


slightly forward. The operculum is of the paleomelanian type, of 
many (6 or 8) turns, and, in the species seen, is considerably smaller 
than the aperture. The perfect apex of the shell (nuclear whorl) is 
asymmetrical. Specimens of the animals of Brotia baccata (Gould) 
and the adult and embryonic shells of three other species of Brotia 
from the Salween River of Thailand and Burma have been personally 
examined. In the species baccata there is a well-developed, deep 
genital groove down the right side of the foot that abruptly turns in- 
ward to become the entrance (or “birth pore’’) of the brood pouch. 
This brood pouch is very large, crowded with a great many relatively 
small young of uniform size. The single character of the embryonic 
shell, or the apex (when perfect) of the adult shell, that characterizes 
Brotia is the asymmetry. It appears as if each shell in its development 
is interfered with by a yolk sac (?) in the position of the apex to keep 
that apex soft, while the rest of the embryonic whorls develop nor- 
mally and symmetrically, and then the apex collapses to a line below 
the curve of spiral symmetry. This may very well be connected with 
a secondary development in the brood pouch to insure a “‘nurse’’ type 
of nutrition of the young. 

Synonyms of Brotia include Acrostoma Brot 1874, Brotella Rovereto 
1899, Paracrostoma Cossman 1900, with the genotype Augeli Philippi, 
and Wanga S. F. Chen (1943, p. 21), based on the species Melania 
henriettae Gray (1834, pl. 18, fig. 2). In this connection it should be 
mentioned that the original figure of henriettae is poorly drawn and 
does not show the generic character of the slightly produced columellar 
angle of the aperture. This fault is corrected in the figure of the 
holotype published by Yen (1942, p. 204, pl. 15, fig. 66). In fact it is 
possible that the originally published locality of henriettae (““China’’) 
is at fault, and this may be the same species as baccata (Gould) of the 
Salween River and some of its tributaries. 


Genus Fijidoma Morrison, 1952 


Genotype: (Fijidoma laddi Morrison 1952)=Fijzidoma maculata 
(Mousson) 1865, by original designation. 

Fiujdoma Morrison (January 1952) antedates by only a few months 
the synonymous generic name Veloplacenta Hubendick (August 1952), 
which was based on the same species of thiarid snail. These fresh- 
water snails from Viti Levu in the Fiji Islands, superficially very 
similar in shell outline to those of the southeast Asiatic genus Paludo- 
mus, were named in allusion to that similarity. They have rather 
smooth, subglobose, neritiform shells with a few rather rapidly in- 
creasing, well-rounded whorls separated by a deep suture. The shell 
has fine spiral sculpture and regular flammules or rows of squarish 
dots (interrupted flammules) of red in the ground color. The aper- 


384 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


EXPLANATION OF PuiateE 11 


1, Melanopsis dufourit (Ferussac), female, from the right side showing egg-laying groove 
and the “ovipositor” in pit on right side of foot, from Elche, Alicante Province, Spain 
(USNM 515845). 

2, Oxytrema canaliculatum undulatum (Say), female, showing egg-laying groove and pit in 
right side of foot, from Kentucky River, Clifton, Ky. (USNM 597434). 

3, Oxytrema nodifila (Martens), female, as in fig. 2, from Han River, Kwang-jang-in, Korea 
(USNM 597451). 

4, Pleurocera zonalis (Rafinesque) (=Melania obovata Say), female, as in fig, 2, from 
Kentucky River, Clifton, Ky. (USNM 595969). 

5, Oxytrema bulbosa (Gould), female, as in fig. 2, from tributary of John Day River, 10 
miles east of Austin, Grant County, Oreg. (USNM 473801). 

6, Paludomus labiosus (Benson) (?), female, as in fig. 2, from roadside run north of Lampang, 
Thailand (USNM 361267). 

7, Oxytrema deshayesiana (Lea), female, as in fig. 2, from Tennessee River, near Florence, 
Ala. (USNM 601564). 

8, Planaxis sulcatus (Born), ovoviviparous and parthenogenetic female showing egg-transfer 
groove and brood-pouch pore (“birth pore”) on side of neck, and subhaemocoelic 
brood pouch in back of head and neck, from Romurrikku Island, Bikini Atoll, Marshall 
Islands (USNM 585106). 

9, Thiara sp., female, showing brood-pouch pore and numerous young in the subhaemocoelic 
brood pouch, from Mindanao, Philippines (USNM 434144). 

10, Brotia baccata (Gould), female, showing deep egg-transfer groove on right side and 
subhaemocoelic brood pouch with hundreds of relatively small young, from Salween 
River, Ban Mae Hick, Burma (USNM 420467). 

11, Thiara sp., female, as in fig. 9, showing the anus, oviduct opening, and brood-pouch 
pore relations, from Naga, Okinawa, Ryukyu Islands (USNM 601311). 

12, Melanoides sp., female, as in fig. 9, with unpigmented shallow “groove,” and numerous, 
equal-sized, small young, from Palo, Leyte, Philippines (USNM 573572). 

13, Tarebia lateritia (Lea), female, as in fig. 9, with numerous small young, from Caminiwit 
Point, Mindoro, Philippines (USNM 487583). 

14, Melanoides turriculus (Lea), female, as in fig. 9, with numerous very small young in 
pouch, from Caminiwit Point, Mindoro, Philippines (USNM 487582). 

15, Melanoides sp., female, as in fig. 9, with a very few proportionately enormous young 
developed in the subhaemocoelic brood pouch, from Manganyama, Rota, Marianas 
Islands (USNM 593472). 

16, Pachychilus (Pilsbrychilus) ‘dalli Pilsbry, old individual showing 3 consecutive peri- 
stomes to prove continuity’ of sinuous lip character of adults, from Tehuantepec, 
México (USNM 133197). 

17, Thiara (Setaeara) cancellata Réding, fresh shell showing epidermal spines, from Siaton, 
Negros Oriental, Philippines (USNM 597436). 

18, Fijidoma maculata (Mousson) (holotype of laddi Morrison) from Viti Levu, Fiji 
Islands (USNM 597433). 

19, Cubaedomus brevis (Orbigny), ovoviviparous and parthenogenetic female (sketched from 
dried material), showing ‘only 2 embryonic young of relatively enormous size in 
subhaemocoelic brood pouch," from Rio los Cayos, Bahia Honda, Pinar del Rio, Cuba 
(USNM 407991). 

(Animal sketches not drawn to scale; figs. 16, 17 slightly reduced; fig. 18 enlarged approxi- 
mately 2 times.) 


U, S, NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 11 


beget AT. 


(FoR EXPLANATION SEE OPPOSITE PAGE) 


OLD AND NEW WORLD MELANIANS—MORRISON 385 


ture is nearly semicircular with a broad, flattened, white columella. 
The operculum is similar to that of Melanoides; thin, corneous, and 
paucispiral, of less than two turns. <A few small-to-medium, but not 
uniform-sized, young shells found dried inside some adult specimens 
indicated that this genus belongs to the Thiaridae, although the 
animals were not available for study. 


Fijidoma maculata (Mousson) 1865. 


Publication of the independent investigation by Hubendick of 
shells and animals in the collections of the Bernice P. Bishop Museum 
has brought to my attention the following synonymy: 

Ampullacera maculata Mousson (1865, p. 203). 

Ampullacera maculata Crosse (1865, p. 432, pl. 14, fig. 10). 
Ampullarina maculata Nevill (1878, p. 249). 

Salinator maculata Hubendick (1945, p. 108). 

Fijidoma laddi Morrison (Jan. 1952, p. 8). 

Veloplacenta maculata Hubendick (Aug. 1952, pp. 179-184, pl. 1). 

In this species the body whorl is enormous, so that the height of 
the spire is less than one-fourth the height of the aperture. The 
aperture is roughly semicircular, with both upper and lower (anterior 
and posterior) angles well rounded. Nuclear,whorls ‘smooth; early 
postnuclear whorls sculptured by incised spiral lines between broad, 
flat rib-areas, crossed by microscopically fine (cuticular) growth lines. 
The spiral sculpture becomes obsolete or absent on the body whorl, 
persisting longest just below the suture and in the columellar region. 
The columella is rather straight, broad, heavily callused, and flattened 
or a little excavated, in contrast to the even curve of the thin outer 
lip. The shells are reddish horn with regular flammules or equivalent 
rows of minute squarish dots of red in the ground color. The broad 
columella is whitish. Embryonic shells of 3 whorls are globular, 
smooth except for growth lines, and even at this early stage possess 
a heavy, callused, but not flattened columella. (See pl. 11, fig. 18.) 

The holotype of F’. laddi (USNM 597433) and a number of paratypes 
(USNM 532559) were collected from rocks in a swift stream, the Lami 
River, on Viti Levu, Fiji Islands, May 30, 1926, by Dr. H. S. Ladd 
of the U. S. Geological Survey. 

The holotype of laddi has 3 whorls remaining, and measures as 
follows: Height, 9.2 mm.; diameter, 8.2 mm.; aperture height, 8.0 
mm.; aperture diameter,5.8mm. The flat, white columella is 1.5 mm. 
wide. 

Comparison of all available figures and specimens has shown that 
there is only one described species of Fizidoma. It has been recorded 
to date from the upstream, swiftly flowing waters of the Rewa River 
and Lami River systems on Vit Levu. 


386 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Hubendick (1952) has rather completely described and figured the 
critical and diagnostic anatomy of animals of this genus. His slight 
misinterpretations of the reproductive anatomy are easily understood 
by anyone who has personally examined the gross reproductive 
anatomy of thousands of individual mollusks belonging to the ‘‘ Mela- 
nian complex,” including hundreds of individuals belonging to the 
family Thiaridae. Nearly all the biological peculiarities seen by 
Hubendick in the seven females of Fizidoma maculata available to 
him are the distinguishing biological characters possessed by every 
individual of the families Planaxidae and Thiaridae (Thorson, 1940, 
and Morrison, 1952). The assumption by Hubendick that these 
animals ever possess a testis must be histologically proven in order to 
refute the observed parthenogenicity of these females. Abbott 
(1952, p. 92) could not find sperm in living material or in histological 
sections of the gonads of six mature adults of the related Tarebia 
granifera (Lamarck). Hubendick did not recognize the completely 
separate, secondary or adventitious nature of the brood pouch of 
these animals. His ‘uterus’ is not a part of the primary gonoduct 
as is the case in the Viviparidae and the Lavigeriinae, hence should 
not be called a uterus. Abbott (1952, p. 92) has pointed out that 
the brood pouch of the Thiaridae is not developed until the animal 
begins to reach maturity. 

Hubendick’s correction of the taxonomic placement of Fijidoma 
was incomplete; his references to its close relationship with the genus 
Emmericia of the family Hydrobiidae were erroneous. Elsewhere 
(Morrison 1949, p. 14) I have put on record the critical and diagnostic 
(male) reproductive characters of the subfamilies of the Hydrobiidae, 
including Emmericia of the Emmericiinae. 


Family Tararmak (in Africa) 
Genus Thiara Reding, 1798 


This typical genus of the family occurs in the east Africa region as 
well as in Asiatic waters. 


Genus Melanoides Olivier, 1804 


The genotype of Melanoides (Olivier, 1804, p. 40), MZ. tuberculata 
(Miiller) 1776, was probably the first member of the family to be 
proved ovoviviparous in the thiarid fashion. Raymond (1852) 
described the brood pouch very well. His tales of the young going 
back into the brood pouch at night, however, must be completely 
discounted. What he did not realize was that there is only a partial 
release of the young at any one time, so that every time he looked 
into the brood pouch of an adult he found young still there—not back 
there again. 


OLD AND NEW WORLD MELANIANS—MORRISON 387 


As mentioned above in the discussion of the genus Potadoma, all 
species of Melanoides are ovoviviparous and parthenogenetic. They 
must also agree with the genotype in the paucispiral character of the 
operculum. The species that differ from this pattern and are incor- 
rectly placed in the genus must be placed elsewhere when their 
anatomic characters become known. 


Genus Tanganyicia Crosse, 1881 


Genotype: (Lithoglyphus rufofilosus EH. A. Smith=) Tanganyicia 
rufofilosa (K. A. Smith) 1880. 

With a shell similar to that of the American genus Cubaedomus, 
Tanganyicia from the shores of Lake Tanganyika differs most notably 
in possessing an operculum that is secondarily concentric around a 
paucispiral center. This African genus Tanganyicia (Crosse, 1881, 
p. 123) has a brood pouch similar if not identical in origin to that 
of all other known genera of the family Thiaridae. The species 
rufofilosa (EK. A. Smith 1880, p. 426) has been anatomically described 
by Moore (1898c, p. 457). The primary reproductive organs are 
identical in their simplicity. The genital groove from the oviduct 
opening to the opening of the brood pouch is of exactly the same type 
(structurally demarcated) as that observed by me in the genus Brotia. 
The brood-pouch pore is on the right side, as in all the other genera. 
Moore (1898c, fig. 3), in his ‘‘semidiagrammatic representation,” 
depicts the brood pouch passing under the oesophageal canal. The 
extensively developed brood pouches of many genera of this family 
may effectively almost surround the pharyngeal or buccal mass, but 
I believe the brood pouch is ordinarily dorsal thereto. Even if the 
brood pouch proves upon complete reexamination only to be beneath 
and expanded to the left of the buccal mass, these differences in 
the subhaemocoelic extent of an identically originating invagination 
can only be further proof of the distinction of the genus Tanganyicia 
from the other members of the family. 

Moore (1898b, p. 307) listed Tanganyicia as a member of the 
Planaxidae, and again (1898¢c, p. 456) stated that rufofilosa, in the 
characters of the radula and alimentary canal, approximates the 
Planaxidae. This statement constitutes unbiased corroboration of 
the present studies which show that the Thiaridae and the Planaxidae 
are relatives from the same stock, since Moore was completely 
unaware of the identical type of reproductive characters possessed 
by the Planaxidae (see Thorson, 1940). 

The statement of Moore that the genital groove is present ‘‘in 
both sexes” may be easily explained by the assumption that he had 
before him nongravid females and gravid females at the beginning 
of a seasonal reproductive period. He mentions only eggs in 


388 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


the brood pouch, so his material must have been collected at the 
beginning of the brood-pouch nursing period for the species. 


Genus Stanleya Bourguignat, 1885 


Genotype: Stanleya neritinoides (E. A. Smith) 1880. 

The figures of this shell look suspiciously like a variant of the 
genus Tanganyicia. The opercular and the animal characters must 
be discovered and compared in order to maintain the separation of 
Stanleya. 


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Preston, Huau B. 
1915. Mollusca, pt. 3 (Freshwater Gastropoda and Pelecypoda), in Blanford, 
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xix-+ 244, 
RAFINESQUE, CONSTANTIN 8S. 
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OLD AND NEW WORLD MELANIANS—MORRISON 393 


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PROCEEDINGS OF THE UNITED3{STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1954 No. 3326 


PRELIMINARY ANALYSIS OF THE FOSSIL VERTEBRATES 
OF THE CANYON FERRY RESERVOIR AREA 


By TuHeopore E. WHITE 


The Canyon Ferry Reservoir will be located on the Missouri River 
below Townsend, the county seat of Broadwater County, Mont. The 
dam is being built about 1 mile downstream from the present Canyon 
Ferry Dam and Power Plant. The maximum pool will inundate an 
area approximately 25 miles long and 3 to 4 miles wide, and will 
include several known fossiliferous localities in Oligocene and Miocene 
sediments. The reservoir area includes one of the very few areas in 
the intermountain basins that have a complete succession of sediments 
from the Lower Oligocene well into the Middle Miocene. Fossil 
faunas are known from other intermountain basins in Montana but 
rarely do they embrace more than one of the time units represented 
here. These faunas are of special interest because they represent an 
environment totally different from that of the White River deposits 
of the plains and offer opportunities to study fossil faunas in the light 
of the principles of climatic zoning which have been worked out for 
recent faunas. 

Most of our knowledge of the Tertiary deposits and their faunas of 
the intermountain basins of Montana is the result of the explorations 
and research of the late Dr. Earl Douglass around the turn of the 
century. His studies have been published principally in the Annals 
of the Carnegie Museum. His explorations have been greatly ex- 
tended and enlarged by Dr. J. LeRoy Kay, of the same institution, 
whose assistance has greatly expedited the work of the River Basin 
Surveys. The studies by Douglass have been further augmented by 
those on the fossil rodents and lagomorphs by Dr. J. J. Burke, who 

273552—54—_1 395 


396 PROCEEDINGS OF THE NATIONAL MUSEUM vou. 108 


was the first to point out that certain genera were common to the 
Oligocene deposits of Eastern Asia and the intermountain basins. 
Also should be mentioned the work of the late Dr. W. D. Matthew on 
the fauna of Pipestone Springs, which is the only study on a single 
fauna which has been made on any of the intermountain basins. 

As part of the salvage program being carried out by the River Basin 
Surveys of the Smithsonian Institution in the reservoir sites in the 
Missouri Valley, the fossiliferous localities in the Canyon Ferry Reser- 
voir area have been prospected for fossils for three seasons. In 1947, 
the first season of work, largely reconnaissance, I was assisted by 
John C. Donohoe, now a student at the University of New Mexico; 
in 1948, by Mr. Donohoe and Ernest L. Lundelius, now a graduate 
student at the University of Chicago; and, in 1950, by William C. 
Harrup, Jr., a student at Columbia University, and Prentiss Shep- 
herd, Jr., a student at Harvard University. With the aid of these 
sharp-eyed young men a collection of nearly 300 specimens was ob- 
tained from the several localities in the reservoir area, the basis of 
thisstudy. Thesespecimensarein the U.S. National Museum(USNM). 
In addition, Dr. Kay has given me free access to the material from the 
reservoir area in the Carnegie Museum (CM) which has materially 
supplemented the collection of the River Basin Surveys. 

The graphic art of William D. Crockett has been invaluable in por- 
traying the characters of the more interesting specimens. 

In the discussion of the specimens, the localities from which they 
were obtained are indicated by code numbers according to the system 
used by the Missouri Valley Project of the Smithsonian River Basin 
Surveys. Following is a list of these localities. 


Lower Oligocene (Chadronian) Localities 


24LC16. NEYSW%, sec. 3, T. 10 N., R. 1 W., of the Montana prime 
meridian, about 1 mile north of Canyon Ferry, Lewis and Clark 
County, Mont. ‘This is a small area of badlands at the foot of the 
bluff on which the buildings of the permanent offices of the project 
are built (fig. 40). About 130 feet of light buff ashy clays are exposed 
at this locality with Chadronian fossils in the upper half. Only the 
remains of the smaller mammals were sufficiently well preserved for 
identification. 

24BW18. NW¥SEX sec. 7, T. 9 N., R. 1 E., of the Montana prime 
meridian, about 14 miles southwest of the south end of Lake Sewell, 
Broadwater County, Mont. This locality (fig. 41) was originally dis- 
covered by Dr. Kay, who very generously directed the River Basin 
Surveys to it. Lithologically, the deposits in this area do not differ 
materially from those of the preceding ‘ocality, nor does the species 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 397 


assemblage. About 300 yards east of this locality some Middle 
Oligocene fossils were collected but were so few that separate locality 
designation was not given. 


Middle Oligocene (Orellan) Localities 


24LC15. SEMSEY sec. 10, T. 10 N., R. 1 W., of the Montana prime 
meridian, about 4 mile southeast of Canyon Ferry, Lewis and Clark 
County, Mont. This is a small butte, about 100 feet high, of light buff 
clayey ash showing indistinct bedding. The greatest concentration of 
fossils is on the south side. However, the fossiliferous area is so lim- 
ited that it very strongly suggests that the fauna was brought together 


oI ees 


PESOS AAO nen 


Ficure 40.—Locality No. 24LC16 from the south. 


by an Oligocene owl. The photograph in the popular account by 
Douglass (1908) shows that this is the Canyon Ferry locality from 
which he obtained his Oligocene fossils. 

24LC17. SWYNEX sec. 3, T. 10 N., R. 1 W., of the Montana prime 
meridian, about 1% miles north of Canyon Ferry, Lewis and Clark 
County, Mont. This locality offers a more complete Middle Oligocene 
section than the preceding one. About 140 feet of sediments are 
exposed at this locality. At the base they are light gray to buff 
clayey ash, grading to nearly pure ash, grayish green, at the top. The 
same faunal assemblage is found in the lower levels as at the preceding 
locality but no identifiable specimens were obtained from the upper 
levels. 


398 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


Lower Miocene (Arikareean) Localities 


24LC18. S¥SW% sec. 11, T. 10 N., R. 1 W., of the Montana prime 
meridian, about 1 mile east of Canyon Ferry, Lewis and Clark County, 
Mont. About 200 feet of fine-grained, dense, buff sandstone, which 
weathers into nearly vertical cliffs are exposed in this area (fig. 42). 
In texture and color these deposits very closely resemble the Harrison 
deposits of western Nebraska and eastern Wyoming. This locality is 
the one from which Douglass and others have obtained Miocene fossils 
at Canyon Ferry and it is still the most productive Miocene locality 
in the area; but, even so, fossils are not common. 

24LC19. NE“%SW% sec. 2, T. 10 N., R. 1 W., of the Montana prime 
meridian, about 1% miles northeast of Canyon Ferry, Lewis and Clark 
County, Mont. This locality is a small area of cliffs in the buff sand- 
stone on the northeast side of the highway just southeast of the bridge 
over Cave Gulch. Very little material was obtained at this locality. 

24L.C20. SWY%NEY sec. 11, T. 10 N., R. 1 W., of the Montana 
prime meridian, about 1 mile east of Canyon Ferry, Lewis and Clark 
County, Mont. This locality lies across Magpie Gulch, about \% mile 
north of locality No. 24L.C18, and represents the lower levels of that 
locality. Only a very small fauna was obtained here. 


Middle Miocene (Hemingfordian) Localities 


24LC21. SEYNW*Y sec. 13, T. 10 N., R. 1 W., of the Montana 
prime meridian, about 2 miles east and little south of Canyon Ferry, 
Lewis and Clark County, Mont. A few isolated and imperfect Mery- 
chippus teeth were obtained from a buff sandstone resting on a lens of 
fresh-water limestone. These few teeth were the only material 
collected in this locality. 

Earl Douglass location. W% sec. 25, T.9 N., R 1 E., of the Montana 
prime meridian, about 8 miles east of Winston, Broadwater County, 
Mont. Douglass (1908b, p. 274) secured the greater portion of a 
skull, a lower jaw, and some skeletal element of an advanced species of 
Merychippus from the bluffs on the east side of the Missouri River. 
And, as far as I know, no other specimens have been collected from 
this locality. 


Geology 


The Canyon Ferry Reservoir area lies in the reentrant between the 
Big Belt Mountains on the east and the Spokane Hills on the west, at 
the north end of the deformational basin in which the Toston beds of 
Douglass (1901, pp. 242-243) were deposited. As near as could be 
determined in the limited time available, the beds are not lithologically 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 399 


continuous with those of the Prickley Pear Valley or the Thompson 
Creek area until at least the Middle Miocene times. Also deposition 
was uninterrupted from Lower Oligocene (Pipestone Springs equiva- 
lent) into the Middle Miocene (possibly Marsland equivalent). In 
Late Miocene times orogenic movements tilted the earlier sediments to 
the northeast and coarse gravels, which were later cemented with 
calcium carbonate, were deposited on their truncated edges. As yet 
no identifiable fossils have been found in these gravels, but they are 
believed to be Pliocene in age. The Pleistocene is represented by 
local deposits of coarse, unconsolidated gravels which have not yet 
produced diagnostic fossils. 


wi 


Ficure 41.—Locality No. 24BW18 from the south. 


The probable conditions under which these sediments were deposited 
are very well described by Douglass (1903, pp. 146-149). Insummary, 
the sediments accumulated in a deformational basin which, in its lowest 
part, harbored a series of small, very shallow lakes or wet meadows 
or a combination of both. Evidence for the lakes lies in the several 
local areas of fresh-water limestones, some of which showgvery fine 
bedding planes. Evidence for the wet meadows lies in the areas of 
dark, gypsiferous clays, often with shalelike partings. However, in 
all probability the gypsum is of secondary development rather than 
primary. 

As pointed out by Douglass (1903, p. 147) the fossils occur most 
abundantly near the margin of the depositional basin and are very 


400 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 104 


rare towards the central portion. One interpretation which could 
be placed on this fact is that only along the margins did sediments 
accumulate fast enough to bury the bone before weathering or other 
factors could destroy them. However, evidence of alluvial fans along 
the basin’s margin have not been observed. The alternative, which 
seems the more probable, is that the accumulating soil of the central 
portion was more acid from decaying vegetation and the bones, react- 
ing with the sulphides of vegetable decay, were a partial source of the 
gypsum in these deposits. 

Throughout their observed exposures the Oligocene sediments are 
uniformly fine grained, ranging in color from light gray to light buff. 


SLA PRIESTS A Og tS REESE RS ty 
rie 


Figure 42.—Locality No. 24LC18 from the south. 


No lithological distinction could be made between the Lower and 
Middle Oligocene sediments, but those which are believed to represent 
the Upper Oligocene contain considerably more ash, are less consoli- 
dated, and grayish green in color. The Oligocene sediments show 
rather even bedding planes which are indistinct except when the ex- 
posures are wet. Occasional stream gravels are found at nearly all 
levels but do not appear to have cut very deep into the Oligocene 
deposits. These gravels are made up almost entirely of angular frag- 
ments of the Proterozoic and Paleozoic sediments which form the 
Big Belt Mountains and the Spokane Hills. 

The Miocene sediments, like the Oligocene, are uniformly fine 
grained but contain a higher percentage of sand and are better con- 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 401 


solidated. In color and texture they very closely resemble the Har- 
rison beds of western Nebraska and eastern Wyoming. Also like the 
Oligocene, the stream gravels occasionally encountered are composed 
largely of the angular fragments of the Proterozoic and Paleozoic 
sediments from the adjacent mountains. Also, basaltic, volcanic 
bombs from 8 inches to 1 foot in diameter are occasionally encountered, 
but they are not necessarily associated with the stream gravels. 
Although the Oligocene and Miocene sediments are quite different 
lithologically, at only one place, in a recent road cut, was the contact 
between the two observed. It was not possible to detect an angular 
unconformity either at this place or by measuring the angle of dip of 
the beds. 

The following section was measured by Mr. Harrup at the north 
end of the reservoir area near the present construction camp; here, 
both the Oligocene and Miocene strata dip east 50° north at an angle 
of 6° from the horizontal: 

Fine-grained, buff sandstone, often weathering into vertical cliffs; stream gravels 
common; occasional voleanic bombs. Arikareean fossils.—150 feet. 

Light gray to buff clayey ash grading to nearly pure ash at top; lenses of stream 
gravels common. Orellan fossils at the bottom.—140 feet. 

Dark gypsiferous clays, with iron nodules, alternating with lighter bands. No 
identifiable fossils—110 feet. 

Light buff clayey ash with Chadronian (Pipestone Springs) fossils at the top.— 
130 feet. 


Class REPTILIA 
Order SQUAMATA 
Suborder SAURIA 

Family ANGUIDAE 


Glyptosaurus cf. montanus Douglass 


USNM 19081, portion of right frontal with scutes and portion 
of left dentary with six teeth, from the Lower Oligocene of locality 
No. 24BW18. 

The limited material of this form does not permit any additions to 
Gilmore’s (1928, p. 115) discussion of the species. 


Peltosaurus sp. 


USNM 19085, a maxillary fragment; and USNM 19084, a dentary 
fragment; both from the Middle Oligocene at Canyon Ferry. These 
specimens appear to be referable to this genus but contribute nothing 
to our knowledge of the group. 


402 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Suborder SERPENTES 
Family Bomar 


Calamagras sp. 


USNM 19082, a single thoracic vertebra from the Lower Oligocene 
of locality No. 24BW18, appears to be referable to this genus. As 
pointed out by Gilmore (1938, p. 36), this genus is difficult to distin- 
guish from the following one, but in view of the very limited state of 
our knowledge it will be convenient to retain one genus for the small 
boids of the Oligocene and the other for the Miocene. 


Ogmophis arenarum Douglass 


USNM 19083, two thoracic vertebrae and the neural arch of a third 
in a connected series, from the Lower Miocene of Canyon Ferry, are 
referred to this species on the basis of the ovate central articulations, 
but the material is too limited to contribute anything to our knowledge 
of the genus. 

It is interesting to note here that of the two other reptilian types 
commonly found in the Oligocene of the plains, one, the crocodilians, 
is unknown and the other, the turtles, is represented only by the 
terrestrial types. No fragments indicative of any of the aquatic 
types have yet been found. On the other hand, the aquatic types 
of turtles are rare in the White River deposits. 


Class MAMMALIA 
Order MARSUPIALIA 
Suborder POLYPROTODONTIA 
Family DIDELPHIDAE 


Peratherium fugax Cope 


USNM 18953, left mandible with P.-M,; USNM 18954, left man- 
dible with P,-M,. Both specimens are from locality No. 24LC15. 

These specimens are nearly twice the size of P. titaneliz Matthew 
from Pipestone Springs and only slightly larger than the measure- 
ments Scott (1941, p. 962) gives for this species. An examination of 
the material in the U. S. National Museum referable to this genus 
shows that the variation in the characters of the teeth and jaws is 
greater in this genus than in the higher mammals. It is probable, as 
Scott points out, that there are only two valid species, a large and a 
small, in the Middle Oligocene of North America. Measurements 


of teeth (in millimeters): 
USNM 18958 USNM 18964 
Le ee ee. SON Seay ks oe A ee. (Se 6. 0 6. 3 
1 AE reeset e MUI Senn ORT eRe Ens Tess (Gul 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 403 


Order INSECTIVORA 


Family SOLENODONTIDAE 
Apternodus mediaevus Matthew 


USNM 18914, fragment of right maxilla with M’, from locality 
No. 24LC16; CM Field No. 30/48, skull and jaws somewhat crushed, 
from locality No. 24BW18. 

The discovery of the skull and jaws was unusually fortunate in 
that they show the character of the enlarged upper and lower first (?) 
incisors. 

Description.—Upper incisoris simple, placed nearly vertical in the pre- 
maxilla, tapering very slightlyin width toward the tip and slanted medi- 
ally so that the tips meet, stronger anteroposterior taper which extends 
from base to tip on the posterior side, anterior face slightly convex 
dorsoventrally and with uniform curvature, enamel covered except 
at tip where worn. Lower incisor is simple, making a very obtuse 
angle with the axis of the jaw, medial face flattened, lateral surface 
convex, very slight taper in width, anteroposterior taper from base 
to tip, tooth enamel covered. 

Family LEPTICTIDAE 


Ictops acutidens Douglass 


USNM 18910, badly broken skull and jaws with skeletal fragments; 
USNM 18912, left maxillary fragment with P*-M*. Both specimens 
are from locality No. 24LC16. 

This material does not permit any additions to be made to Mat- 
thew’s (1903, p. 207) discussion of the species. 


Family TALPIDAE 


Genus and species undetermined 


USNM 18915, right humerus lacking the distal epiphysis, from 
locality No. 24L.C016; USNM 19024, right humerus lacking the distal 
epiphysis, from Pipestone Springs; CM 9184, right humerus lacking 
the distal epiphysis, from locality No. 24BW18. 

Discussion.—A comparison of this material with the humeri of 
recent moles in the U. S. National Museum does not reveal any 
close affinities with any of the genera represented. On the other 
hand, the derivation of the humerus of the Talpinae and Scalopinae 
from this type of humerus would require only an exaggeration of its 
present characters. Consequently, the reference of this material to 
the Talpidae is reasonably certain, but generic designation will be 
withheld pending the acquisition of better material. 

273552542 


404 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


The discovery of this material confirms the suggestion (Simpson, 
1947, p. 637) that this group probably migrated to North America 
from Eurasia during the Early Oligocene. Also of interest is that 
the group reached North America before the humerus acquired the 
extreme modification which it has today. 


Family NycriTHERIIDAE 
Kentrogomphios strophensis, new genus, new species 


Genoholotype—USNM _ 18870 (fig. 43), facial portion of a skull 
anterior to the cribiform plate, lacking one canine, the crown of the 
other, and the incisors. 

Referred material ——_USNM 18871 (fig. 44), right mandibular frag- 
ment with Mz-_3. 

Horizon and locality —lLower Oligocene, Chadronian; SW% sec. 3, 
T. 10 N., R.1 W., of the Montana prime meridian, about 1 mile north 
of Canyon Ferry, Lewis and Clark County, Mont. 

Diagnosis.—Portion of skull preserved about the same size as the 
corresponding portion of Scalopus aquaticus; skull elements securely 
fused together, orbit small, infraorbital foramen large and opening 
into a deep elongate pit anterior to orbit, lacrymal duct large and 
situated within.the orbit, no palatine vacuities, zygoma believed 
present but very slender, foramina of the alisphenoid region similar 
to those of the Soricidae; I?, C1, P3?, M3; root of canine elongate- 
triangular in cross section with the acute angle posterior, cusps of 
cheek teeth very high, posterior border of P* to M? deeply emarginate 
between hypocone and metastyle, no anterior or inner cingulum, 
strong cingulum on hypocone; P? (if present) minute, single rooted 
and simple; P* three rooted, a single outer cusp with a broad shearing 
blade extending posteriorly, deuterocone minute and placed nearer 
the anterior than the posterior edge; P* submolariform, parametacone 
twice as high as protocone, strong metaconal crest, parastyle and 
metastyle strong, external cingulum weak, protocone twice as high 
as hypocone, strong posterior cingulum on hypocone; M! with para- 
cone and metacone close together and subequal in height, metastylar 
crest strong, parastyle and metastyle well developed, mesostyle ob- 
solete, a small metastylule present, protocone strong and as high 
as paracone, a small protoconule present, hypocone half as high as 
protocone and with a strong posterior cingulum; M? similar to M?’, 
paracone and metacone farther apart, a strong parastylar crest, 
parastyle and metastyle well developed, parastylule and metastylule 
present, a strong mesostyle present; M? triangular in outline, hypocone 
and metastyle absent, parastyle and parastylule present, a minute 
metaconule present, mesostyle well developed. 


VERTEBRATES—WHITE 405 


CANYON FERRY FOSSIL 


os Wy = 
V ne ee 

ee 

iy 

a ys 

1 

ras 

if 

i 


USNM 
18870 


Figure 43.—Lateral and occlusal views of the type (USNM 18870) of Kentrogomphios 


Strophensis, new genus and species.  X 7. 


406 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


Referred specuomen.—Trigonid on M, high, height twice that of the 
talonid and nearly twice the length of the tooth, recurved and com- 
pressed anteroposteriorly, protoconid slightly higher than paraconid and 
metaconid, which are equal, an anterior cingulum present which 
terminates in a small but distinct cusp about the middle of the base of 


ee 


Se USNM 
———— 188 7/ 


Ficure 44.—Occlusal and lateral views of the referred specimen (USNM 18871) of Kentro- 
gomphios strophensis, new genus and species. X 7. 


the paraconid; talonid nearly as long as the trigonid but narrower, 
hypoconid high and with a ridge sloping down to the base of the 
metaconid, height of hypoconid nearly equal to the length of the tooth, 
hypoconulid minute, entoconid nearly obsolete; M; similar to M, but 
smaller and with a narrower talonid. 

The portion of the jaw preserved indicates a strong upward curva- 
ture back of the teeth. The massateric fossa is deep and with a very 
prominent anterior rim. 

This specimen is referred to this species on the basis of the occlusion 
with the type which is as good as occurs between different specimens of 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 407 


the same species. The two specimens are from the same horizon and 
locality but the difference in amount of wear excludes them from 
belonging to the same individual. 

Discussion.—Although this specimen adds many new data to our 
knowledge of this group of insectivores, the affinities of the family are 
still problematical. A number of the characters are shared by both 
the Soricoidea and the Microchiroptera. Some characters are shared 
by some members of both groups but not by all members of either. 
Briefly, these characters are the fused skull elements and the char- 
acter of the orbits. This is also true for the general characters of the 
cheek teeth. In some members of both groups the posterior border 
is deeply emarginate between the hypocone and metastyle. The 
parastylar and metastylar crests resemble both groups in their general 
features but in their details resemble the bats more than the shrews. 
The very high cusps of the cheek teeth, especially the hypocone, 
appear to be a unique feature of this specimen. The large canine 
excludes this specimen from theshrews but not from the moles (Talpinae) 
and bats. The elongate pit in front of the orbit is a characteristic of 
most shrews and many of the moles, but not of the bats. A slender 
zygoma is present in moles and most bats but is absent in the shrews. 
The rostrum is broken away and the anterior termination of the face 
cannot be determined. 

In summary, the characters of this specimen exclude it from the 
Soricidae but do not definitely affiliate it with any other group. The 
molelike facial features certainly suggest that it could belong to the 
talpid humerus from the same locality. Parascaptor and Scapato- 
chirus have a large canine and reduced premolars as does this specimen. 
However, the cheek teeth of this specimen are of a type unknown in 
any living moles and very suggestive of the bats. Measurements of 
teeth (in millimeters) : 


USNM 18870 
Cayenne +. OF Sadi hha VaR Joie Jot Panh. Lars 11. 6 
po aaa SRE a Greedy eho At. ee Bie 3 OL. Nis da St 3. 6 
1 a Be Se ere eee eee eee ee eee 5.5 
Length Width 
Ee ee Da RG, | ee Dae ie 
ieee Sere Oe ee Tk SEN eS ee Sa ee ee Re 8: 2a 
MiG A ode Pa oho OER UE ved py ek ed 1.8 PA N7/ 
OLS Bae 2 A ten 2 aa Mee TES EAS PEM Fees 2s ae 1B PL? 
USNM 18871 
Width Height 
Length Trigonid Heel Trigonid Heel 
MM, BOY iS 7 90d 20 01d WS CES 2 Shik es 
Mg. 22 etiaricianie tad) SHO 0.8 Dil 1.0 
Depth of jaw between M, and M3_-_-_-___-- PAS 


Depth of jaw below masseteric fossa_ -_ -___- 1,2 


408 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Order RODENTIA 
Family ScruRAVIDAE 
Prosciurus cf. relictus (Cope) 


USNM 18857, left mandible with P,-M;; USNM 18858, left man- 
dible with P,-M;. Both specimens are from locality No. 24LC15. 

The enamel patterns of the teeth and the measurements agree 
with the figures and description by A. E. Wood (1937, p. 168) but 
this material does not permit any additions to his discussion. Meas- 
urements of teeth (in millimeters) : 


USNM 18857 
Length Width 
| EIN a eee Sap ae aes es Spey Re ee. 1. 2. See 
| Ei. een a, it A, eek Seine eich erica Mats nT 15 1. 6 
AVE; SERRE AR AS Sb SE nee eR) Se BNE) IL) Dare aE ee 1.6 ile? 
NiL@ J.) POLO S10) BOT PO a iG ey Ee Od MOT! ie DOF, 1.9 
Miavtnacode » os pidan gdb toy tana pe je ata 2. 2 1. 6 


Family [scoyROMYIDAE 
Ischyromys cf. pliacus Troxell 


USNM_ 18908, right mandible with P,M;; USNM 18909, left 
mandible with P<M,; USNM 18920, right mandible with M,-Ms3. 
Specimens are from locality No. 24LC16. 

This material does not permit additions to our knowledge of the 
species. 


Titanotheriomys veterior Matthew 


USNM 18904, a palate with dP;-M;; USNM 18905, a palate with 
dP.-M,; USNM 18907, left mandible with P,-M;. Specimens are 
from locality No. 24LC16. 

This material does not permit any additions to our knowledge of 
this species. 


Family CAsToRIDAE 
Paleocastor cf. gradatus (Cope) 


USNM 18963, palate with left P*-M°, from locality No. 24LC18; 
USNM 18962, left mandibular fragment with P,—Mz, from locality 
No. 24LC20. 

This material is referred to this species on the basis of size. As 
pointed out by Stirton (1935, p. 409), this species is very close to 
P. peninsulatus (Cope). Definite allocation of the Canyon Ferry 
material must await the acquisition of better specimens. 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 409 


Family EomyipAE 


Paradjidaumo cf. minor (Douglass) 


USNM 18759, left mandible with P;—Msz, from locality No. 24LC16; 
USNM 18758, right mandible with P,-Mg, from locality No. 24LC17. 

Discussion.—In size, this material agrees with both size groups 
which Wood (1937, p. 244) records from Pipestone Springs, but it 
differs in that M, and M, are broader than long in both specimens. 
One cf the specimens measured by Wood (AMNH 9635) shows this 
condition, but with such a limited sample it is impossible to properly 
evaluate this character. 

Although one of these specimens is from the Chadron equivalent 
and the other from the Lower Brule equivalent, I could find no 
morphological characters of the teeth which would separate them. 
Also, in the Paradjidaumo material from Pipestone Springs in the 
U. S. National Museum there is one specimen which agrees with 
P. trilophus in its dental morphology, including the flattened anterior 
face of the lower incisor. In view of the above facts and the variation 
in size and tooth proportions found in the material from Pipestone 
Springs and the Badlands, it is difficult to escape the inference that 
P. minor and P. trilophus represent species groups rather than single 
species. Measurements of teeth (in millimeters): 


USNM USNM 

18758 18759 
Length Width Length Width 
J a a aa aaa eer eee Oe ee eee 
pees Bees en oak ee LA fo, £2 eo 
LM cece — pape pS CN EN pe del on BPD! AIG = aires 1. 4 
i. BURL Dota 8 GO BOleDI97_£ Se WSLS 1.4 
Me mre are 6 qt het menos 28h L Gai -betlasreeiose 42. 


Paradjidaumo spokanensis, new species 


Holotype —USNM 18760, a badly crushed skull with right mandible. 

Horizon and locality Middle Oligocene, Orellan; SEYSE¥ sec. 10, 
T. 10 N., R. 1 W., of the Montana prime meridian, about 4 mile 
southeast of Canyon Ferry, Lewis and Clark County, Mont. 

Diagnosis.—Upper teeth broader than long with mesoloph extending 
to labial margin, posterior cingulum uniting with hypocone and 
extending nearly to labial margin, mure extending nearly to middle of 
tooth, shortest on P* and longest on M?. Lower teeth with mesolophid 
long, extending to lingual margin, free on P, and M, but uniting with 
entoconid on M, and M; with wear. P, longer than broad and Mz; 
nearly as broad as long. Measurements of teeth (in millimeters): 


410 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Upper Lower 

Po-My_......-2- 32.5 SESE 6. 0 627 
Length Width Length Width 
Py. - 2 - = 2 22 | (OR AOI ae IS63PAIN7> 1.7 1.5 
pe eed Ae ee i res 15°: 1.6 
MG =. 20 UU BOOL BADGES Behr eh PY Hy eas GAGE LNG Ua 
Megs. 0.41 Var enok mr? alAa "> Lin 1S ZBL ITS 1.4 


The characters which distinguish this species are the larger size, the 
elongate P,, and the short, nearly quadrangular M3. 


Family CRICETIDAE 


This family of rodents is represented in this area by a single genus, 
Eumys, and most of the specimens came from two localities, Nos. 
24LC15 and 24LC17. The others, specimens in the Carnegie Muse- 
um, are from the upper levels of No. 24BW18, locally known as the 
Old Hadcock Ranch. This series of nearly 100 specimens, although 
showing considerable variation, is divisible into four groups, three of 
which exhibit characters of the teeth not readily referable to species 
described from the White River deposits of the plains. All of the 
specimens are 15 to 25 percent larger than those of the plains. Also, 
the teeth are higher crowned and the cusps are better developed. 

By analogy with some of the wide-ranging species of the genus 
Peromyscus, one would normally expect only subspecific differences 
between the members of the genus Humys from the Badlands of South 
Dakota and the Canyon Ferry Reservoir area since the former is on 
the plains and the latter is in the mountains. On the other hand, in 
the mountain areas two or three distinct species of Peromyscus 
usually occur in a single area in addition to the plains species. This 
appears to be correlated with the greater variety of ecological niches 
within a limited area, and the remains of a varied fauna from a 
variety of ecological niches could be concentrated in a very small area 
of deposition by the work of owJs. In view of the geologic history of 
this region, the environment for small rodents during the Oligocene 
was probably as varied and as different from that of the plains as it 
is today. Although only one of the plains species has been positively 
identified in this material, A. E. Wood (1937, p. 250) reports LZ. 
elegans Leidy from Montana but gives no locality data. 

The following artificial key gives the distinguishing characters 
of the species of this area: 


Key to the species of Eumys 


a.1 Anteroconid on M; large, anterior cingulum on Mg, extending the full width 
of tooth, a short but distinct mesolophid on M»____-_-- cricetodontoides 
a.2 Anteroconid on M; small, mesolophid on Mz, obsolete or absent. 
b.1 Cross lophs normal. 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 411 


c.1 Lingual portion of anterior cingulum on Mz, about half as long as 


A sela el Soper iigs Sk pe Bh rd oe a SR Oh Si latidens 

c.2 Lingual portion of anterior cingulum on M; obsolete or absent. 
spokanensis 
b.2 Cross lophs weak, ectolophid strong__..........-..------------ exiguus 


Eumys cricetodontoides, new species 


Holotype —USNM 18748 (fig. 45), right mandible with the incisor 
and M,-3. 

Horizon and locality.—Middle Oligocene, Orellan; SEY%SE¥, sec. 10, 
T. 10 N., R. 1 W., of the Montana prime meridian, about \% mile 


Figure 45.—Occlusal and lateral views of the type (USNM 18747) of Eumys cricetodontoides 
new species. X 7, 


Referred material—_USNM 18749-18754, six mandibles from the 
same locality. 

Description of type-—M, elongate, anteroconid large, nearly as 
wide as tooth, anterior arm of protoconid united with anteroconid 
and posterior arm with the metaconid, mesolophid short but distinct, 
posterior cingulum not united with entoconid; M, quadrangular, 
longer than broad, anterior cingulum extending across anterior 
border of tooth, lingual and labial portions subequal, posterior arm 
of protoconid extending nearly to lingual margin of tooth, closely 
applied but not united with metaconid, mesolophid short but very 


273552—54——_3 


412 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


distinct, posterior cingulum not uniting with entoconid; M, subtrian- 
gular in outline, sharply constricted behind protoconid, lingual portion 
of anterior cingulum short but distinct, posterior arm of protoconid 
extending to lingual border of tooth and united with metaconid at 
base, no mesolophid. 

Variation.—In one specimen the anterior arm of the metaconid is 
united to the anteroconid while the protoconid is not. In one, both 
are united to the anteroconid. The mesolophid of M, and Mz is 
variable in length but is always distinct. In two specimens the 
posterior arm of the protoconid M, and M; is shorter than in the type, 
but the teeth agree in other particulars. 

The short mesolophid on M, is very suggestive of Cricetodon, but 
the evidence does not warrant referring this material to that genus. 
Measurements of teeth (in millimeters): 


2 M Ma Ms 

USNM No. < Length Width Length Width Length Width 
LS749-4 Aes See eerie 8.0 PA, 1.9 2. 6 222) e2a0 2.0 
TS pees Se eee 2 oe ee 8. 2 2.9 2. 2 2. 6 2. 4 25 yar 
1S Clipe) o'=- 2 te See oe GS 22.5 1.9 2. 8 2. 2 2. 5 21 
WSyoeek 22k Soe See Sees = aco 2.8 1. 8 2.5 2.2 ee 2. 0 
IS {beet 2 Se ee coe eee 7.9 2.8 2.1 2. 6 2. 2 2. 6 2.1 
1S 7508. See eee eee Soe eee oa 2. 6 1. 2.3 2. 0 2. 6 2. 0 
1S 7 gee wee ees a oe BIC see. 6h ero i eae eee 2.1 


Eumys latidens, new species 


Holotype-—USNM 18772 (fig. 46), right mandible of young individ- 
ual with M,-3. 

Horizon and locality.—Middle Oligocene, Orellan; SEYSEY sec. 10, 
T. 10 N., R. 1 W., of the Montana prime meridian, about % mile 
southeast of Canyon Ferry, Lewis and Clark County, Mont. 

Referred material—USNM _ 18764-71, 18773-80, 16 mandibles 
from the same locality. 

Description of type-—M, elongate, anteroconid small, protoconid 
united to anteroconid and metaconid, mesolophid well developed and 
distinct, posterior cingulum not united with entoconid; M, quad- 
rangular, longer than wide, anterior cingulum extending across front 
of the tooth, lingual portion shorter than the labial, both protoconid 
and mesoconid united to anterior cingulum, posterior arm of proto- 
conid not united to metaconid and not extending to lingual border 
of tooth, mesolophid obsolete, posterior cingulum not uniting with 
entoconid; M; sharply constricted behind protoconid, lingual end 
of anterior cingulum obsolete, posterior arm of protoconid approaches 
but does not unite with metaconid, no mesolophid. 

Variation.—The material assigned to this species is extremely 
variable in many of the dental characters but the designation of 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 413 


another species does not seem warranted at this time. The antero- 
conid of M; is always small but the anterior cingulum is variable 
in extent. In one specimen, both the protoconid and metaconid are 
united to the anteroconid. This variation occurs in all three species 
but is not consistently associated with other characters. The 
mesolophid is variable in extent but is always distinct. On Mz, the 
lingual portion of the anterior cingulum is variable in extent but is 
always distinct. The posterior arm of the protoconid varies in 
length but never extends to the inner border of the tooth. The 
mesolophid is consistently obsolete or absent. On M; the lingual 
portion of the anterior cingulum may extend nearly to the border of 


Ficure 46.—Occlusal view of the type (USNM 18772) of Eumys latidens, new species, 
anterior end to the right. X 7. 


the tooth or may be nearly obsolete. The posterior arm of the 
protoconid may or may not unite with the metaconid. Measurements 
of teeth (in millimeters): 


M, M3 M3 

USNM No. Mit ngth Width Length Width Length Wiath 
eye Uiype) 2S So et Fear 2. 6 LOW V2S5 20 2H 2S203 222 
Leyp( As SRE) pe eee eae eee ee Soy 248 1.9 PAREN pees) 247 232, 
LC Pieg a ote gee td eee a oe 7.9 2. 6 Devil 2. 6 2. 4 2.5 2.3 
SYS a ee ee GA oC 24 WO ey ee De Dr 2.41 
DIAPER ho be ee Tae 2. 6 1.9 2. 4 22 7A! ot 
Lil. 2 ido! alge ert helen eile Me WO a ae CoO ee Pas Vel oy Acs yA 
Livi Bo te AD 8. 2 23 pe | 26 pa Det 2.3 
LE? Ti. | ig eS ee (Ga) PASTS 1.9 2. 4 253 2a Dal 
L/S eee WO 2. OD £2 Og CBO se Dass 24 22 2 
Ly ee rs ee Moe 255 19 2; 4. | 2. 4 ee | 
L200 h(E ee ar eee ee ar ee 8. 1 Zon. v2.0) 2.6. 2.0 2.50 2.0 


Eumys spokanensis, new species 


Holotype —USNM 18833 (fig. 47), right mandible with M,-3. 

Horizon and locality Middle Oligocene, Orellan; SE¥SEY sec. 10, 
T. 10 N., R. 1 W., of the Montana prime meridian, about % mile 
southeast of Canyon Ferry, Lewis and Clark County, Mont. 


414 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Referred material—Eleven mandibles from the same locality and 
C. M. 9136 from the upper levels of locality No. 24BW18, locally 
known as the Old Hadcock Ranch. 

Description —M, elongate, anteroconid small, anterior cingulum 
well developed, anterior arm of protoconid connected to anteroconid 
and posterior arm to metaconid, mesolophid well developed and ex- 
tending nearly halfway to inner margin of tooth, inner end of posterior 
cingulum close to but not united with entoconid; M; quadrangular, 
nearly as broad as long, lingual portion of anterior cingulum obsolete, 
posterior arm of protoconid closely applied to entoconid and extend- 
ing nearly to inner margin of tooth, no mesolophid, posterior cmgulum 
as in M,; Mg; subtriangular in outline, sharply constricted posterior 
to protoconid, anterior cingulum as in Mg, posterior arm of protoconid 
united to metaconid at base, no mesolophid or mesoconid. 


Ficure 47.—Occlusal view of the type (USNM 18833) of Eumys spokanensts, new species, 
anterior end to the right. X 7. 


Variation.—This species shows less variation than the other two. 
The teeth have the appearance of being shorter, for their breadth, 
and the cusps more crowded, but the measurements do not bear this 
out. On M,; and M; the posterior arm of the protoconid often be- 
comes united with the metaconid with wear. The lingual portion 
of the anterior cingulum of M, and Ms; is represented by a vertical 
fold near the midline of the tooth and soon becomes obliterated with 
wear. Measurements of teeth (in millimeters) : 


Mi M2 M3 

USNM No. Mrs Tcagih Width Length Width Length ava 
Tessa CL yDe)eee oa eee St (2e7 1.9 -254.--* 22 2a ieee 2.0 
TS Ss Fie eas os) See es 7.4 25 1.8 2.5 2a PRS 2.0 
TSSAS aby. ee eae NA kas re eee 7.8 12.6 2.09) U285 PAE 2.4 2.0 
MSSAQ iis Se Ee. gh ee a Te ees HENS 220 1.9 DA all 2.5 2.0 
USS41 ole =) ak ee he a 8.1 2.9 22026 2.25 2.3 aa: Poe 
WSS47 os cee eee ee 2 TATE DIF 2.0 74.03" a2, 23 2.1 
FLSS OSS) esere ee ee tee iw! EE Rene ap qa 2.6 1.9 2. 4 742 2. 4 2.0 
LSSAGe AVN Beers % 8°-"2.6 19 4203. * "242-4 2nG 2.3 
$8842.20... 3Oe oY sti ie 8. 0 Ha 1.9 yA 222 2. 4 2. 2 
PSSA5 Leer tae ee ee A et SESE we 7.6 PATI 18. 4222 2-0) - 25 2.0 


1 Estimated. 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 415 


Eumys cf. exiguus Wood 


USNM 18866 and 18867, 2 mandibular fragments with M)-3; 
USNM 18865, 1 maxillary fragment with M,-;. All specimens are 
from locality No. 24LC17. 

This material is provisionally referred to this species on the basis 
of the tooth cusp arrangement. While these specimens are larger 
than the type, the sample is too small to furnish grounds for separate 
designation. Measurements of teeth (in millimeters) : 


Mi M: M3; 
USNM No. Mit J ongh Width Length Width Length Width 
SU ee ee ere ty SPS GA Pil UG he iL. 7 A203 iL} 7¢ 
MSOs ee io 2s. ree h. et ft 6. 4 IB} 1.4 2. 1 1.5 1.8 1.5 
HSRC ee ee ee es ls 247, 13,9),.4 2:0 HEH Ae eee Ke 
Order LAGOMORPHA 


Family LEPoRIDAE 


Paleolagus temnodon Douglass 


USNM 18869, right maxilla with P*-M*; USNM 18875, right 
maxilla with P?-M?; USNM 18876, right maxilla with P?-P*; USNM 
18877, right mandible with P;-M;. Specimens are from locality Nos. 
24LC16 and 24BW18. 

This material does not permit any additions to Wood’s (1940, p. 
320) discussion of the species. 


Paleolagus intermedius Matthew 


USNM 18872, left mandible with P;-M;; USNM 18873, left man- 
dible with P,-M.; USNM 18874, right maxilla with P?@-M?. Speci- 
mens are from locality Nos. 24LC15 and 24LC17. 

This species appears to be relatively rare in these deposits and the 
limited material does not permit a satisfactory comparision with the 
specimens from the plains. 


Paleolagus burkei Wood 


USNM 18879-18894, 4 upper and 12 lower dentitions. All of the 
specimens are from locality Nos. 24LC15 and 24LC17. 

The upper dentitions are slightly larger than the measurements 
given for the type (Wood, 1940, p. 327), but they agree quite closely 
in the details of the tooth form and do not agree with P. haydeni 
Leidy. I have not seen any material from the Middle Oligocene of 
this area referable to the latter species. It is possible that this 
species could have been restricted to the mountains and P. haydeni 
Leidy restricted to the plains during the Middle Oligocene and that 
P. burkei Wood did not spread to the plains until the Upper Oligocene. 


416 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Megalagus brachyodon (Matthew) 


USNM 18903, right mandible with P,-M;,;; USNM 18902, right 
mandible with P;-M,. Both specimens are from locality No. 24LC16. 

This limited material does not permit any additions to Burke’s 
(1936, p. 150) discussion of this species. 


Archaeolagus sp. 


USNM 19096, right maxillary fragment with P*-M?, from locality 
No. 24LC18. 

This specimen is so incomplete that the generic reference is only 
provisional. The teeth are greatly flattened, the transverse diameter 
is twice the anteroposterior. The hypostria is deep, extending nearly 
to the middle of the tooth, a little closer to the posterior edge than 
the anterior, and extending to the base of the tooth. Groove on the 
lateral surface of the tooth shallow and broad. No crescentic valley 
(this may be a state of wear). No enamel on the lateral surface of the 
tooth. Cement present only in the hypostria (this may be an accident 
of preservation). The teeth appear to be rootless. 


Order CARNIVORA 
Family CANIDAE 


Hesperocyon paterculus (Matthew) 


USNM 18911, skull and jaws (fig. 48) with the greater portion of 
the skeleton; USNM 18897, right mandibular fragment with M,_3; 
USNM 18896, left mandibular fragment with M, 4 2; USNM 18895, 
left mandibular fragment with P, . 3, Mi-3. Specimens are from 
locality Nos. 24LC16 and 24BW18. 

Description.—Dental formula: I?, Ci, Pi, M3. Upper dentition: 
P' single rooted and closer to C than to P?; P? with anterior cingulum. 
incipient posterior cusp and minute cusp on heel; P*® with well developed 
anterior cingulum, posterior cusp and a cusp on heel; P* with incipient 
anterior cusp; M! without incipient metaconule, paracone higher than 
metacone, protocone well developed, protoconule small but distinct, 
hypocone well developed, anterior cingulum terminates at base of 
protocone; M? similar to M! but smaller; M* minute and probably of 
no systematic significance. Lower dentition: P; 42 missing, P; 
single rooted and closer to C than to P.; P; 4 4 differ only in size, each 
with well-developed anterior basal and posterior cusp, heel well 
developed and with a small cusp; M; with well-developed hypoconid 
and incipient entoconid; M, with protoconid and metaconid sub- 
equal and elevated above the paraconid, hypoconid well developed, 
entoconid incipient; M3 small, protoconid and metaconid subequal. 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 417 


Discussion.—The general form of the skull of this species does not 
appear to be noticeably different from the other small dogs of the 
Oligocene of the plains, nor is it possible to detect differences in the 
skeletal material. The characters of the teeth of these specimens 


USNM 
I89/! 


Ficure 48.—Upper and lower dentition of Hesperocyon paterculus Matthew (USNM 
18911). X 1. 


418 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


appear to be quite constant and agree very well with the material 
from Pipestone Springs in the U. S. National Museum. The variety 
of tooth-cusp arrangement and height of crown exhibited by the 
small carnivores of the Middle Oligocene which have been referred 
to H. gregarius (Cope), both in the literature and in collections, make 
any attempt at comparison with that species entirely futile. However, 
these specimens are easily distinguished from the single specimen of 
Hesperocyon from the Lower Brule equivalent of this area. 

The specimens of small carnivores of the Oligocene of the plains 
which I have examined in several museums convince me that the 
species represented by this tooth-cusp arrangement had a very close 


Figure 49.—Occlusal view of the first and second lower molars (USNM 18898) of Hespero- 
cyon gregarius (Cope) showing dental caries. X 4. 


relative (if not the same species) in the Middle Oligocene. But until 

the material has been carefully studied and the indeterminate types 

have been relegated to nomena nuda we are helpless taxonomically. 
Measurements of teeth (in millimeters) : 


Length Width 

TSI VIS So ON ee 33. 0 
i 5 ag Ree aa Ra Re ig ee Ay sd 24. 5 
"i Nog: phapea eave Seis salut amu Rene Pures IF, «5 16. 0 

Anterior Posterior 
IVE aN Olt ek Me Ee 6. 5 7. 5 7.0 
Manse 3 es ae bn ee ese NE 3. 5 5. 0 4.5 
MB a A eh EB eal 1.0 1:25, 2es2=2 

Trigonid Heel 
Be Mig ccs ce ee i Be ge: 34.0. 
Pe = ak fe ern oe Ree 20..0. 3 eee 
Van Ss: «SA ey eee eee: hg 9. 0 4.0 3. 5 
Mig. Di > Sea ee hs bee BAL eS 5.0 3. 5 3. 0 
M3222 seat oeeus’ . yt oe eee ee 2. 0 2. 0:28.22 


Hesperocyon gregarius (Cope) 
USNM 18898 (fig. 49), right mandibular fragment with M,_», from 
locality No. 24LC15. 
The specimen referred to this species differs from the preceding one 
in that on M, the entoconid is well developed and about half the size 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 419 


of the hypoconid. A metaconulid and a protoconulid are present as 
in Nothocyon and Tomarctos. There is a small but distinct entoconid 
on Mz. 

An unusual feature of this specimen is that both teeth show evidence 
of dental caries. On M, there is a major lesion on the hypoconid 
(fig. 49) with a minor lesion on the entoconid, protoconid, metaconid 
and metaconulid. On M, there is a major lesion on the protoconid 
with a minor lesion on the metaconid and hypoconid. Among the 
recent carnivores the only evidence of dental caries I have seen was 
in the teeth of the grizzly bears (Ursus horribilis Ord) in the Museum 
of Comparative Zoology and in the U.S. National Museum. Meas- 
urements of teeth (in millimeters): 


Width 
Length ~ oe 
Trigonid Heel 
iho = Sie Se es ee a a 11.0 5. 0 4.0 
Vigna ee nk es ee i 2 AM 5. 0 4.0 3.5 


Nothocyon cf. geismarianus (Cope) 


USNM 19097, right mandible with P,—-M,, from locality No. 
24LC19. 

This specimen is slightly smaller than the one described by Cope 
(1884, p. 920) but agrees in other details. 


Family UrsipaE 


Subfamily AMPHICYNODONTINAE 
Daphoenocyon cf. dodgei (Scott) 


USNM 19094, left mandibular fragment (fig. 50) with dP3_, and 
M,-2, from locality No. 24LC16. 

This specimen is badly fractured but it is possible to make out the 
essential details of the teeth. The deciduous teeth are more canidlike 
in the cusp arrangement than are the permanent teeth, but are low 
crowned. IndP; the anterior and posterior cusps are well developed, 
the posterior accessory cusp present and distinct, heel broad with pro- 
nounced internal and external cinguli. In dP, the trigonid does not 
differ from that of the permanent tooth except in size, hypoconid and 
and entoconid subequal with entoconid slightly larger, a distinct 
hypoconulid slightly smaller than the hypoconid present. The per- 
manent teeth, being in an unworn condition, exhibit features on the 
heels of M, and M, which appear to be common to many members of 
this subfamily, namely, in the breaking up of the principal conids into 
smaller cusps. In this specimen, while the conids on the heel of M, 
are quite distinct, the hypoconid has two apices and the entoconid 
has three. The heel of M; is a further exaggeration of this phenome- 


420 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


non. The hypoconid and entoconid are represented by two ridges 
which are beset with small tubercles and meet posteriorly. 

Hough (1948, p. 594), noting the striking difference between the 
type of this species and the other species of Daphoenus, proposed a 
new genus, Daphoenocyon, to receive it. At the same time she erected 
a new family, the Daphoenidae, to receive a number of North American 
genera which had been included in the Caninae but which, quite 
obviously, had little in common with the true dogs. On the other 
hand, there are three groups, the Amphicynodontinae, the Amphi- 
cyoninae, and the Simocyoninae—all of holarctic distribution from 
the Lower Oligocene to the Pleistocene and well established in the 
literature—into one of which any member of the new family could be 
fitted without changing the definition of the group. Also, to include 
the Simocyoninae, as represented by Protemnocyon, in the same family 
with the other two subfamilies is an unnatural and nongenetic 
grouping. 

Since the characters of the auditory region (Hough, 1948, p. 577) 
and the post cranial skeleton, insofar as it is known, of the Amphi- 
cynodontinae and the Amphicyoninae are closer to the bears than the 
true dogs, it is my opinion, in view of the evolutionary fate of the 
former, that these two groups should be placed in the Ursidae rather 
than being placed in a separate family or included in the Canidae. 
That the early members of these two subfamilies should possess a 
generalized canoid dentition is to be expected if they are to be in- 
cluded in that superfamily, but it is a fallacy to use the superfamily 
characters of the teeth to determine the family and subfamily affini- 
ties when these determinations contradict the affinities shown by the 
fundamental structure of the limbs and basicranium. 

A comparison of the figures of D. dodge: (Scott) and Parictis 
dakotensis Clark (Scott and Jepsen, 1936, pl. 12, fig. 3; pl. 14, fig. 1) 
shows that these two genera are indeed closely related and belong 
to the same subfamily. This relationship was confirmed by the com- 
parison of a series of jaws in the Carnegie Museum from Pipestone 
Springs which are referable to these two genera. In fact, except for 
very minor details, the only difference between the two is size. In 
both forms the teeth are low crowned, rather broad, and distinctly 
less bladelike than in the more typical canids. Although, in the 
material available, size is the only character which distinguishes the 
two genera, it is probable that when the material is better known 
valid distinctions will be found. 

Although specimens of this subfamily of carnivores seem to be 
relatively rare in North America, I suspect that the scarcity is more 
apparent than real and that quite a number have been referred to 
either Hesperocyon or Daphoenus in the collections of our various 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 421 


museums. I have prepared table 1 (below), partly from the 
literature and partly from my knowledge of the collections in a few 
museums, to show the distribution of this group in North America. 
An examination of this table shows that the smaller form, Parictis, 
persisted from the Lower Oligocene into the Lower Miocene, while 
Daphoenocyon continued only into the Middle Oligocene (Hough, 1948, 
p. 595) and we have no record of it after that. In the Lower Miocene 
two new forms appeared, Pachycynodon and the form from Florida 
which I described as Parictis bathygenus White (1947, p. 500). Even 
though the available data are limited, they appear to substantiate 
the opinion (Simpson, 1947, p. 630) that this group is Eurasian in 
origin and immigrated to North America. Also it appears that there 


SG: a 
Pr... 
» 


Ficure 50.—Occlusal view of dP3-4 and Mi-2 (USNM 19094) of Daphoenoeyon dodgei 
(Scott), xX 1%. 


was a migration in the Lower Oligocene and another in the Lower 
Miocene, since neither Pachycynodon nor Parictis bathygenus are 
at all closely related to the American Oligocene forms. 


TABLE 1.—Distribution of the Amphicynodontinae in the Early Tertiary of North 


America 
Locality Chadronian Orellan Whitneyan Arikereean 
Pipestone Springs Parictis 
Daphoenocyon 
Canyon Ferry Daphoenocyon 
John Day Parictis 
Wyoming, Nebraska, and South Parictis Parictis Parictis Pachycynodon 
Dakota Daphoenocyon Daphoenocyon 
(Hough, 1948) 
Florida Parictis bathygenus 
Order PERISSODACTYLA 
Family EquipaE 


Mesohippus hypostylus Osborn 


CM 9184, left maxilla with M*-*; CM 8998, palate with right and 
left P?-M?; unnumbered, right maxilla with M'-*; USNM 18946, skull 


422 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


with teeth badly worn; USNM 18949, right mandible with P.-M;; 
CM 9171, right mandible with P;-M.; CM 9365, right mandible with 
P.-M.; CM 9363, right mandible with dP:_.; CM 9364, right mandible 
with dP,_,. Specimens are from locality Nos. 24LC16 and 24BW18. 

Discussion. —In size and tooth characters the material agrees better 
with this species than any of the many which have been described 
from the Lower Oligocene. Also, the small crochet, which character- 
izes M. portentus Douglass, is lacking. 

Since this material is more complete than any previously recorded 
from this area, it will be described briefly. 

Description.—Upper dentition: P? with protocone well developed, 
protoloph weak, protoconule indistinct, metaloph well developed with 
distinct metaconule, hypostyle distinct, mesostyle weak but distinct, 
parastyle distinct, no metastyle; P’, P*, and M! quadrangular, pro- 
toconule and metaconule distinct, parastyle and hypostyle distinct, 
metastyle weak, internal cingulum between protocone and hypocone, 
mesostyle strong; M? similar to M! except metaconule very indistinct; 
M! similar to M? except much narrower posteriorly. Measurements 
(in millimeters): 


CM 8998 CM 9184 
P2-M?2_____ GONO Vee ton mie th teteet-e ee 
Bc Foe ee 3020) Ubware,, aietn ett oes 
14 6 Ss RO ee ee eee RT ee mM ae ts 40. 2 
Height at Height at 
Length Width metacone Length Width ‘metacone 
| FW Mea Sa ali ia ot 1D. S34 ol223 sissy 2. 22 Mg JOG), Gee 
P38. 2h ee ED OS9T4NOY? Siege prs. J lA! betes eee 
Pees Se 2 ee eee 12,0, WosOW sees ose oe noon oso 
Mi Aeebteye 2 ee 13.5 15.0 (worn) 8.0 13.7 15.8 10. 0 
1 a a 12.8 16.0 85" 1454. 1652 8. 6 
IMS Lae Abie at te | ge AAS. EER eee 13.0) 15. bee 


Lower dentition: P, subtriangular in outline, protoconid reduced, 
metaconid not twinned, entostylid small but distinct, strong external 
cingulum; P;, Ps, M,, and Msg essentially similar, strong external, 
anterior, and posterior cinguli, small median external cusp, metaconid 
and entoconid distinctly twinned, parastylid indistinct; M,; with meta- 
conid indistinctly twinned and entoconid not twinned. Measurements 
(in millimeters): 


CM 9171 CM 9365 

Pg Mga Sees ee re ee ee 64 
Pye Miser era sacs ee re ae 52 52 

Length Width Length Width 
Paha Livics 22 atR s emesitorry. hee fut. ebos. eliDades aSaG 
Pas deme eet Ss ce an gee bere ne 13..0 | 1009, --= ae 
] 5) PINS Ra 4 as Reale Oe ase 13.0 “10:6 ~13.07 Ie 2 
1 (ae SP SS ARE ene A ie tt SE Ne See Eas 12.6 9.0 12.4 10.0 


Pet ee ker ee emt 8 Oar P a oT ot os a 127 68.3 AZ 6 1s 


* Locality of Earl Douglass 


Co 


‘ = 


Y 


S 
eek 


Insects -Leaves 


a Fossiliferous Localities 


FROM BUREAU OF RECLAMATION MAP 
BS/IS/ S51 


Salar 273552 O- 54 
localities. 


=—"™~ 


NORTHERN PORTION OF THE CANYON FERRY RESERVOIR AREA 


c- 


Avalanche 
Gulch 


Hellgate 


Gulch 
Jc 
“ 
¢ 
_— CO 
ww Fossiliferous Localities 
Lake Sewel/ 
Missouri River Sy 
@ 
y 2 FROM GUREAU OF RECLAMATION MAP 
= a/ 15/51 
“=~ Ne 
: VS 


Fieure 51.—Northern portion of Canyon Ferry Reservoir area showing location of the fossiliferous localities. 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 493 
Merychippus sp. 


USNM 19095, two fragmentary lower molars, from locality No. 
24LC21. These teeth are too imperfect for specific identification but 
are sufficient to establish the presence of Middle Miocene deposits in 
this area. 


Superfamily BRONTOTHERIOIDEA 


At locality No. 24LC16 a few fragments of teeth referable to the 
large members of this group have been found but they are inadequate 
for even generic reference. 


Family HyracoDONTIDAE 


USNM 19025, right mandibular fragment with dP;—M,, from locality 
No. 24BW18. 

Discussion.—This material is too fragmentary to permit allocation 
to any of the species described from the White River deposits. Sin- 
clair (1922, pp. 65-79) recognized four species on the characters of the 
upper premolars and stated (p. 67) that no intermediate stages were 
recognized. Scott (1941, p. 841) expressed the opinion that there was 
only one valid species of this genus in the White River deposits on 
the grounds that four species of a single genus of large mammals could 
not occupy the same territory equivalent in size to that embraced 
by the White River deposits. On a previous page (p. 786) Scott 
quotes Matthew’s (1930, pp. 271-272) views in connection with the 
species of Trigonias. In his introduction Matthew states that: “We 
do not, in fact, find two or more distinct species or subspecies of a 
genus occupying the same area and habitat at the same time.”’ This 
statement appears to have been accepted by some workers as an axiom 
but, stead of inspiring caution and the use of recent faunas in inter- 
preting fossil faunas, it has been used as a legitimate excuse for multi- 
plying the number of genera. With the aid of Dr. Henry Setzer of 
the Division of Mammals, U. S. National Museum, I have prepared 
u short list of instances where two or more distinct species of the 
larger mammals occur in the same area and habitat: 

CrRvIDAE.—Within historic times the ranges of the Virginia deer (Odocoileus 
virginianus) and the mule deer (O. hemionus) overlapped by nearly a million 
square miles and they occupied the same habitat over the area. No natural 
crosses are known. ‘These forms are morphologically distinct and the skulls and 
horns would be recognizable as fossils. 

CanipaE.—The above paragraph is true of the wolf (Canis nutilus) and the 
coyote (C. latrans). 


Equipar.—The horse (Equus caballus) and the ass (EH. asinus) ran wild in 
western North America for about 300 years after escaping from the Spaniards 


424 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


and no natural crosses are known. Also £. prezevalskit and E. onager occur to- 
gether in Central Asia in the wild state and no undisputed cases of crossing are 
known. 

BovipaE.—In the Malay region Lydekker (1898) records four species of Bos 
(gaurus, indicus, depressicornis, and frontalis) occurring in the same area. 

ANTILOPIDAE.—Allen (1939) reports five species of Gazella (dorcas, leptoceros, 
thomsonii, granti, and soemmerringit) occurring together in the Anglo-Egyptian 
Sudan. 

Even with this short list it is obvious that two or more species of a 
single genus of the larger mammals do occur together over large areas 
and in essentially the same habitat, and by analogy it is entirely pos- 
sible that more than one species of a genus could be found in a “quarry 
fauna.” 

In his treatment of the fauna of a formation, Matthew (1930, p. 272) 
points out that there is a time factor involved and that shifts in ranges 
(possibly due to minor climatic fluctuations or seasonal movements of 
the herds) would make it possible for species with adjacent ranges to be 
found as fossils in a single time unit of aformation. This is illustrated 
by the ranges of the barren-ground caribou (Rangifer arctrcus) and the 
woodland caribou (2. caribou), both of which range across Canada 
from east to west with a very narrow overlap in range. Even though 
the amount of overlap did not change, a minor climatic change would 
permit one to occupy a greater amount of the former territory of the 
other and the two might be found together as fossils, dependent, of 
course, on the accidents of preservation. 

Another method by which species of adjacent areas may be intro- 
duced into an area of deposition is by floods. I think anyone who has 
ever seen the bloated carcasses of cattle floating in the streams of the 
west during a summer flood will admit that such a possibility cannot 
be overlooked in regard to the fluvial deposits of the plains. Even if 
this happened only once in each climatic microcycle (11 years) it 
would be sufficient to establish a species in a single time unit of a for- 
mation. It is obvious, of course, that the ‘‘visiting’’ species must. 
have its range upstream fron the area of deposition. A shift in the 
range of extraterritorial forms might be the explanation of the pres- 
ence of Caenopus in only the Chadron and Upper Brule and its ab- 
sence in the Lower Brule. 

Another item which must be kept in mind in the consideration of 
fossil faunas is the territorial range of the individual. Some of the 
large predators, such as the mountain lion (Felis concolor) and the 
grizzly bear (Ursus horribilis), have a territorial range with a 200-mile 
radius, while with the large herbivores it seldom exceeds 50 or 60 
miles. In the case of small mammals, such as Microtus, the radius of 
its territorial range seldom exceeds 20 or 25 feet. Even with this 
limited number of examples it is clear that with the same food habits 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 425 


(carnivore or herbivore) the probability of dealing with a purely local 
fauna is in inverse ratio to the size. 

This is not an attempt to justify the multitudinous species of 
Mesohippus or the subspecies of Trigonias osborni but to point out, 
as did Matthew, that we must utilize the principles of distribution 
and ecology, which have been worked out by the neozoologists, in in- 
terpreting the fossil faunas. 


Family RHINOCERATIDAE 


Caenopus cf. mitis (Cope) 


USNM 19026, right mandibular fragment with P.-M3;, from 
locality No. 24BW18. 

This specimen is referred to this genus and species on the measure- 
ments of the teeth which agree with those of the type. The teeth 
of this specimen are well worn and it may not be identifiable. 


Order ARTIODACTYLA 


Family LEPTOCHOERIDAE 
Leptochoerus sp. 


USNM 18919, right mandibular fragment with P,-M3;, from Lower 
Oligocene of locality No. 24LC16. 

The teeth on this specimen are so badly worn that only provisional 
reference is possible. However, it is sufficient to record the presence 
of this group in these deposits. 


Family MERYCOIDODONTIDAE 
Merycoidodon? sp. 


Only maxillary and mandibular fragments were obtained from the 
Oligocene deposits and these are inadequate for certain generic 
reference. The size of these specimens indicate an animal of about 
the size of Merycoidodon culbertsoni (Leidy). Nothing in the size 
range of the small and poorly known genera has yet been found. 


Mesoreodon chelonyx Scott 


USNM 19091, skull, jaws and greater part of the skeleton; USNM 
19092, skull and jaws; both from the Lower Miocene of locality 
No. 24LC18. 

These specimens are slightly larger than those recorded by Schultz 
and Falkenbach (1949, p. 154) from the same locality, but are still 
within the limits of individual variation. 


426 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


Promerycochoerus montanus (Cope) 


USNM. 19089, skull and jaws with rostrum missing, from Lower 
Miocene of locality No. 24LC18. 

This specimen is that of an old individual with the teeth well worn. 
However, the specimen is only slightly crushed and the size and con- 
figuration can be determined with reasonable certainty. 


Cyclopidius simus Cope 


USNM 19086, skull and jaws, from the Lower Miocene of locality 
No. 24L018; USNM 19088, skull and jaws, from same locality; 
USN. 19087, right mandible with dP;—M). 

Discussion.—Thorpe (1937, p. 242) lists six species of Cyclopidius 
from the Miocene deposits of the Smith River area and concludes that 
only one species (C. simus Cope) is valid. Koerner (1940, pp. 856- 
858) describes two new species from this area, but he does not use 
the same set of characters to distinguish the two species from each 
other or from C. simus. Nor does he present a standard set of 
measurements for the three species. Consequently it is impossible 
to appraise the validity of his species from his treatise. 

The best preserved specimen is somewhat larger than the measure- 
ments which Thorpe (1937, p. 291) gives for C. simus or its synonyms. 
It is nearly as large as C. lullianus Thorpe, but the details of the facial 
vacuities are very different from that species and agree with C. semus. 


Family CAMELIDAE 


Poébrotherium cf. eximium Hay 


USNM 18944, right mandibular fragment with P;-M,; USNM 
18945, left mandibular fragment with dP.-,; CM 9301, right man- 
dibular fragment with P,M,. All specimens are from locality 
No. 24BW18. 

This material is too fragmentary for accurate specific determina- 
tion but is adequate to establish the presence of this genus in these 
deposits. 


Family HyPERTRAGULIDAE 


Leptomeryx transmontanus Douglass 
Leptomeryx? esulcatus Matthew, 1903. Bull. Amer. Mus. Nat. Hist., vol. 19, p. 222, 
fig. 15. 

USNM 18931, 18932, 18934-18939, eight mandibular fragments, 
and USNM 18933, one maxillary fragment, all from locality No. 
24LC16; USNM 18940-18943 and CM 9293, five mandibular frag- 
ments, and CM 9291, 9304, two maxillary fragments, all from locality 
No. 24BW18. 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 427 


Frick (1937, p. 625) synonomized L.? esulcatus Matthew (1903, 
p. 222, fig. 15) with L. transmontanus Douglass. On the following 
page he indicates that the specimens referred by Matthew (1903, p. 
224) to L. mammifer Cope should be considered a large variant of 
this species, and on page 629 he refers the specimens figured by 
Matthew as L.? esulcatus Cope to L. evansi Leidy. The upper denti- 
tions of Leptomeryz in the U. S. National Museum and in the Carnegie 
Museum from Pipestone Springs and Lower Oligocene of Canyon 
Ferry agree with the figure and published measurements given by 
Douglass (1903, p. 167, fig. 11) for the type of L. transmontanus. 
Also, a comparison of this material with the type (USNM 157) and 
referred material of ZL. evansi Leidy from the Middle Oligocene reveal 
a number of differences which are fairly constant and are presented 
in table 2. As far as it was possible to do so the characters of 
L. evansi were taken from the type. 


TaRLe 2.—Contrasting characters of Leptomeryx evansi Leidy and 
L. transmontanus Douglass. 


Character L. transmontanus L. evansi 
Median internal cusp on M1 weak or absent strong 
Median internal cusp on M3 weak or absent strong 
Median internal cusp on Ms small strong 
Postero-internal cingulum on Ps strong weak 
Posterior ridges on P3 inner always connects with outer connects with heel 
heel, outer rarely 
“Paleomeryx fold’”’ on lower molars absent present and uniting with pos- 
terior crescent with wear 
Mesostyle on M1 strong absent 
Mesostyle on M2 weak absent 
Mesostyle on M3 weak absent 
Anterior cingulum on lower molars weak or absent strong but soon obliterated by 


wear 


Matthew (1903, p. 223) stressed the differences in the length of the 
posterior ridges on the protoconid of the third lower premolar as the 
distinguishing characteristic of this species and it is valid for the 
majority of the specimens I have examined. Normally, there are an 
inner and an outer cusp on the heel of P3, each witha short ridge ex- 
tending antero-medially (axis of the tooth). These ridges usually 
meet and fuse near the middle of the heel. Normally, the inner ridge 
from the protoconid is united with this common meeting point. In 
the material at my disposal the outer ridge from the protoconid is 
variable in length, is usually bifid, and in one specimen it joins the 
heel at the common meeting point mentioned above. Occasionally 
the ridge from one of the cusps on the heel does not develop, or the 
ridge may extend directly anteriorly. In two specimens the ridges 
of both heel cusps have grown anteriorly and met the posterior ridges 
from the protoconid, giving P; the appearance of P,. 


Leptomeryx evansi Leidy 


USNM 18924-18926, three maxillary fragments; USNM 18923, 
18927-18930, five mandibular fragments; and seven unnumbered 


428 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


mandibular fragments in Carnegie Museum; all from locality No. 
24LC15; and CM 8995, two mandibular fragments, from the upper 
levels of locality No. 24BW18. 

A comparison of this material with the type of LZ. evansi Leidy 
(USNM 157) and the abundant material from the Middle Oligocene 
of Wyoming fails to reveal any constant differences in the details of 
the upper and lower cheek teeth. This material averages about 
3 percent smaller than the type, but the sample is too limited for this 
to be of any significance. 


Climatic zoning of Lower Oligocene faunas 


The possibility of climatic zoning of fossil faunas has received very 
little consideration in the literature. This is partially due to the lack 
of data (widely separated fossiliferous deposits of equivalent ages 
have become known only recently) and partially to the rather general 
belief that the Early Tertiary climates were uniformly mild. In 
regard to the latter, Berry (1922) pointed out that while the over-all 
climate of the Eocene may have been much milder than today the 
“arctic flora’’ of that time was not only distinct from that of temperate 
latitudes but was of circumpolar distribution. He also points out 
(1922, p. 13) that the southern limit of the Eocene “arctic flora’’ is 
about 15° north of the southern limit of the existing species of the 
same genera and that the present isotherm (p. 9) would have to swing 
15° to 20° northward to permit the present existence of the same 
floras in the same areas from which we know the Eocene floras. Such 
a change would only eliminate the Arctic Zone of C. Hart Merriam 
as it is defined today and would by no means be sufficient change to 
produce a uniformly mild warm-temperate to subtropical climate. 
Simpson (1947, pp. 645-654) briefly cites evidence for climatic zoning 
in the Miocene in Asia and considers that climate was probably a 
relatively important selective factor in intercontinental migration. 
He also points out (p. 652) that while the evidence for climatic zoning 
is meager there is none against it. 

The possibility of climatic zoning in the Tertiary was first brought to 
my attention by the relative abundance of the protoceratids in the 
Miocene of Texas and their extreme scarcity in equivalent deposits 
of Nebraska and Wyoming. These animals are large enough that 
they would not be easily overlooked. There are few areas in North 
America which have remained as consistently productive over a 
period of years as has the Miocene of Nebraska and Wyoming. Yet, 
from this area, Syndyoceros is known from only two specimens and 
Prosynthetoceros is entirely unknown. Even on such meager evidence 
it seemed highly probable that the factors which controlled the 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 429 


present distribution of mammals in North America had been in 
operation throughout the Tertiary. The climatic zoning of mam- 
malian distribution in North America by C. Hart Merriam (1892) is 
probably familiar to nearly everyone and will not be further discussed 
here. Many workers have greatly elaborated and refined the original 
statement (with some adverse criticism), but the basic concept is 
still valid. 

It is the purpose of the present study to compare the Lower Oligo- 
cene faunas of the plains and the intermountain basins of Montana 
in the light of possible climatic zoning. 

The statement often has been made that the nondiscovery of a 
particular form in a given deposit is not proof that the form did not 
live in that area while the sediments in question were being deposited. 
In general this is true, but to accept it as an axiom is to exclude from 
consideration all problems of distribution and intercontinental migra- 
tion (see Simpson, 1947, p. 652). Probably the greatest value of the 
statement is that it is a very effective intellectual counterbalance 
against overenthusiastic speculation. For application to a specific 
problem it must be weighted against the answers to a number of 
questions: 

1. Was the area accessible to the animal in question? 
2. Is it known from older or younger deposits from this or closely adjacent 
areas? 
3. Is it known from deposits of equivalent age from adjacent areas? Or, more 
remote areas? 
4. If so, how frequently is it encountered? 
5. Is it associated with other genera belonging to the same family? 
6. How frequently are the related genera encountered? 
7. Are the related genera found in the deposit in question? 
8. Are the probable habits of the animal in question such that its remains 
would have a good chance of being buried and preserved? 
9. How extensively and thoroughly have the deposits in question been explored? 
10. Was the climate and environment of the area in question essentially the same 
as that of the adjacent area? Of the more remote area? 
11. What was the size of the animal? 
12. What was the probable territorial range of the individual estimated from the 
size and inferred habits? 


If the faunas of Pipestone Springs and Canyon Ferry (see table 3) 
are compared in the light of these questions it will be seen that, since 
the two areas are so close both geographically and ecologically, any 
faunal differences must be attributed to the accidents of preservation 
and discovery. 

On the other hand, Pipestone Springs and the Badlands of South 
Dakota are separated by a distance of 600 miles (air line), 3° of lati- 
tude, and 3,000 feet of altitude, and both have been collected exten- 
sively. Although the climate of Lower Oligocene times may have 


430 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


been warmer as a whole, the environmental contrasts between the two 
areas must have been as great then as they are now, and it is permissible 
to expect these contrasts to find as much expression in faunal differ- 
ences, particularly among the small, nonvolant mammals, as they do 
today. A comparison of the Lower Oligocene faunas of the two areas 
(table 3) shows that faunal differences did exist. By analogy with 
Recent faunas the differences appear to be correlated with environ- 
mental differences. It is expected that further explorations will 
modify the faunal differences somewhat; but, in view of the extensive 
explorations in both areas, radical changes are not anticipated. 


TARLE 3.— Distribution of Lower Oligocene fauna 


Pipe- Can- Pipe- Can- 
stone yon stone yon 

Species Plains Springs Ferry Species Plains Springs Ferry 
Peratherium titanelix sp. >< sp. Hoplophoneus robustus x 
Apternodus midiaevus x x Mesohippus celer x 
Apternodus altitalonidus x Mesohippus portentus x x 
Micropternodus borealis x sp. Mesohippus latidens x 
Talpa? sp. x x Mesohippus montanus x 
Clinopternodus gracilis x Mesohippus hypostylus x x 
Metacodon magnus x Hyracodon ef. acridens x sp. 
Kentrogomphios strophensis x Trigonias osborni x 
Domnina thompsoni x Caenopus mitis x sp. 
Ictops dakotensis x Colodon occidentalis x 
Ictops acutidens x Colodon cingulatus x 
Ictops thompsoni, ete. x x Titanotheres many sp. sp. 
Prosciurus vetustus x x spp. 
Prosciurus jeffersoni x Aepinacodon americanus x 
Ischyromys pliacus x x Archaeotherium cf. crassum x 
Titanotheriomys veterior x x x Archaeotherium marshi x 
Cylindrodon fontis x x Archaeotherium scotti x 
Pseudocylindrodon neglectus x Archaeotherium mortoni x 
Eutypomys cf. thompsoni x Perchoerus ef. nanus x 
Adjidaumo minimus x % Perchoerus minor x 
Paradjidaumo minor x x x Bothriodon americanus x 
Paleolagus temnodon x x x Heptacodon sp. x 
Megalagus brachyodon x x Merycoidodonts x x x 
Megalagus turgidus x Merycoidodon affinis x 
Desmatolagus dicei x Bathygenys alfa x 
Hyaenodon montanus x Limnetes sp. x 
Hyaenodon cruentus x Hypertragulus chadronensis x 
Pseudopterodon minutus x Hypertragulus crawforden- 
Daphoenocyon dodgei a x sis x 
Parictis dakotensis x Leptomeryx transmontanus x x x 
Hesperocyon paterculus x x Leptomeryx annectens x 
Hesperocyon gregarius Hypisodus paululus = 
Plesictis priscus x Bere ae: x 

j Pseudoprotoceraslonginaris x 
Paleogale inflex x Eotylopus reedi = 
Deinictis ef. fortis 2s Poebrotherium eximium x x x 
Hoplophoneus o’harrai x Stibarus montanus sp. x 
Hoplophoneus mentalis x Leptochoerus or Stibarus sp. x 


For purposes of discussion, the fauna of the Mountain Province as 
here used is the combined faunas of Pipestone Springs and the Lower 
Oligocene of Canyon Ferry. These two areas were chosen deliberately 
because they are separated from the plains by more than one range of 
mountains and cannot possibly be interpreted as having been in the 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 431 


Transition Zone as can the deposits in Weld County, Colo., and the less- 
well-known faunas of Bates Hole and Beaver Divide, Wyo. Thomp- 
son’s Creek, McCarty’s Mountain, and the Drummond Beds have 
been omitted because they have been incompletely reported upon, 
both faunistically and stratigraphically. 

The fauna of the Plains Province has been compiled from Scott, et al. 
(1936-41), Clark (1937), Cook and Cook (1933), and Cook (1934), 
This compilation of the Plains Province fauna may be introducing 
error into the comparisons because, with a north-south extent of nearly 
600 miles for the Oligocene deposits of the plains, latitudinal zoning 
may have existed. But, with so few detailed studies of limited areas, 
definite evidence either for or against latitudinal zoning is wholly 
lacking. 

The comparison of these two faunas is limited by necessity to the 
smaller mammals because the remains of the larger forms, particularly 
the titanotheres and rhinoceri, are often too fragmentary for more 
than family identification. However, such fragmentary evidence is 
sufficient to establish the group in the area in question. Those cases 
where the generic identification is reasonably certain but no specific 
identification can be made are indicated in table 3 by the abbreviation 
“sp.” In the following paragraphs an attempt is made to evaluate 
the differences found in the two faunas. 

InsEectivora.—Although representatives of this group are not 
numerous in collections, one finds it difficult to escape the inference 
that it is due to their small size. There are two genera, Apternodus 
and Jciops, common to both areas but the species are distinct. Two 
genera, Clinopternodus and Metacodon, appear to be confined to the 
Plains Province and, as yet, are unknown from younger deposits 
outside of the plains. Two genera, Micropternodus and Domnina, 
appear to be confined to the Mountain Province. Micropternodus 
does not appear to have survived anywhere beyond the Lower Oli- 
gocene, and Domnina is well represented in the Plains Province in the 
Middle Oligocene. Consequently, one would expect to find it in the 
Lower Oligocene of the plains. On the other hand, the alternative 
that it did not spread to the plains until Middle Oligocene times (as 
appears to be the case with some other genera) is entirely within the 
limits of possibility. Since Kentrogomphios and Talpa? are known 
from single specimens they are useless for this study. 

RovEentia.—Prosciurus is common to both areas in both the Lower 
and Middle Oligocene. JIJschyromys, a large active form, appears to be 
restricted to the Mountain Province in the Lower Oligocene but is well 
represented in the Middle Oligocene of the plains, where it underwent 
considerable radiation. Titanotheriomys and Cylindrodon are common 
to both areas but did not survive into the Middle Oligocene. Pseudo- 


432 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


cylindrodon appears to be restricted to the mountains in the Lower 
Oligocene but did not survive into the Middle Oligocene. Hutypomys, 
a large active form, is common to both Provinces in the Lower Oli- 
gocene. Both genera of the Eomyidae are common to both 
Provinces in both the Lower and Middle Oligocene. 

LacomorpHa.—Megalagus and Paleolagus are common to both 
areas. The species are distinct in Megalagus and probably also in 
Paleolagus. Desmatolagus, as with Domnina and Ischyromys, appears 
to be restricted to the mountains in the Lower Oligocene but is repre- 
sented in the Middle Oligocene of the plains. 

Carnivora.—The creodonts are rare in the Lower Oligocene of both 
Provinces and consequently are useless for this type of study. 
Daphoenocyon and Parictis are common to both areas and no specific 
separation appears possible. Hesperocyon is common to both 
Provinces but the species are distinct. Plesictis appears to be restric- 
ted to the Plains Province and Paleogale appears to be restricted to 
the mountains in the Lower Oligocene but is well represented on the 
plains in the Middle Oligocene. To the best of my knowledge, not 
even fragmentary evidence of the felids has been found in any of the 
Lower Oligocene deposits of the intermountain basins. With such 
complete negative evidence for an area as extensively collected as 
Pipestone Springs, it seems reasonable to infer that the mountains 
were climatically unsuited to the Lower Oligocene felids. 

PERISSODACTYLA.—Mesohippus is common to both Provinces but 
the Lower Oligocene species have not been reviewed since the time 
it was considered an act of the greatest discourtesy to place 
someone else’s species insynonomy. Consequently the data furnished 
by the species of this genus are not suitable for this study. The 
remains of the Rhinocerotidae from the Mountain Province are 
fragmentary but Hyracodon and Caenopus were common to both 
Provinces. The remains of the Brontotheroidea are too fragmentary 
for generic identification. Colodon is common to both areas but its 
scarcity makes it unsuited for this study. 

ArtTIopAcTyLA.—Although the Leptochoeridae are common to both 
areas, their remains are too rare to be suitable for this study. The 
Entelodontidae appear to be restricted to the plains, and, like the 
felids, seem to have found the mountains climatically mhospitable. 
The Tayussuidae are unknown in the Mountain Province but are also 
rare in the plains and consequently unsuited for this study. Likewise, 
the Bothriodontidae are unknown in the mountains and are rare on 
the plains. The remains of the Merycoidodontidae in the Mountain 
Province are very fragmentary and useless for this type of study. Of 
the Hypertragulidae, only one genus, Leptomeryz, is common to the 
two areas but no specific separation between the two regions can be 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 433 


made. Hypertragulus and Hypisodus appear to be restricted to the 
plains and, like the felids, may have found the mountains climatically 
unsuitable. Heteromeryx is known from a single specimen and there- 
fore unsuited for this study. The same is true for Pseudoprotoceras. 
Of the Camelidae, Hotylopus is known from only three or four specimens 
and consequently is unsuitable for this study. Poébrotherium appears 
to be common to both Provinces but the remains in the mountain 
area are very fragmentary and consequently the genus is unsuitable 
for this study. 


TaBLE 4.— Distribution of genera in the Lower and Middle Oligocene 


Lower Middle Lower Middle 
Oligocene Oligocene Oligocene Oligocene 
Moun- Moun- Moun- Moun- 
tain Plains tain Plains tain Plains tain Plains 
Genus Prov. Prov. Prov. Prov. Genus Prov. Prov. Prov. Prov, 

Domnina* x x Brontotheriidae x x 
Ischyromys x x Apternodus x x x x 
Desmatolagus* x x Ictops x x x x 
Paleogale* x x Prosciurus x x x x 
Metacodon x x Eutypomys x x x x 
Plesictis* x x Adjidaumo ve x x x 
Deinictis* x x Paradjidaumo x x x x 
Hoplophoneus x x Megalagus x x x x 
Archeotherium x x Paleolagus x x x x 
Perchoerus x x Daphoenocyon* x x x x 
Bothriodon* x a Parictis* x x x x 
Heptacodon* x x Hesperocyon ve x x x 
Hypertragulus x x Mesohippus x x x x 
Hypisodus x x Colodon x x x x 
Micropternodus x Rhinocerotidae x x x x 
Pseudocylindrodon x Leptochoeridae x x x x 
Clinopternodus x Merycoidodontidae x x x x 
Cylindrodon x x Leptomeryx x x x x 
Titanotheriomys x x Poébrotherium x x x x 


*Subfamilies to which these genera belong are believed to be immigrants to North America from Eurasia 
in Early Oligocene times (Simpson, 1947). 


Table 4 reveals some very striking contrasts between the faunas of 
the two Provinces, particularly in the number of genera in the Plains 
Province which are not yet known from the intermountain basins. 
Most of these forms are medium to large in size and the subfamily, 
at least, would be recognizable on very fragmentary evidence. Con- 
sequently their nondiscovery at Pipestone Springs indicates, if not 
complete absence, that only occasional stragglers entered the area 
during cycles of maximum abundance. Four of these genera, embrac- 
ing three subfamilies, are believed to be immigrants from Eurasia. 
One of the subfamilies, the Anthracotherinae, with its hippopotamus- 
like habits, quite obviously would have found the smaller, swifter moun- 
tain streams entirely unsuited to its way of life. In the case of Plesictis 
and Deinictis (if this is the true situation), it would appear that a 
temperature factor was involved. Of the nonmigrant genera, one 
family, the Hypertragulidae, is of special interest as only one genus, 
Leptomeryz, appears to have been able to invade the mountains and 


434 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


is present in fair abundance. In view of the relative scarcity of 
Hypertragulus, Hypisodus, and Heteromeryx in the Middle Oligocene 
compared to Leptomeryz and in view of the Late Tertiary development 
of this suborder, one finds very attractive the suggestion that the 
center of development and dispersal was more southern and central 
and that the first three genera were very near the northern limit of 
their range. 

That none of the genera of the Plains Province invaded the Moun- 
tain Province between the Lower and Middle Oligocene and the rather 
long list of genera common to the two Provinces during both ages 
indicate that the indigenous genera had become adjusted to their 
environment, or ecologically stabilized, by the beginning of the 
Oligocene and that there were no major climatic changes before the 
close of the Middle Oligocene. 

The Eurasian immigrants embrace six subfamilies. One, the Am- 
phycynodontinae, quickly became adjusted to both Provinces; two, 
the Nimravinae and Anthracotherinae, were restricted to the plains; 
one, the Mustelinae, was divided between the two areas; and two, the 
Soricinae and Ochotoninae, appeared torequire a period of readjustment 
before invading the plains. The apparent absence of Domnina in the 
Lower Oligocene of the plains is certainly open to question since its 
very small size greatly reduces its chances of discovery. On the other 
hand, the zonal distribution of mammals is more clearly reflected by 
the small species, which is possibly a result of the much smaller terri- 
torial range of the individual. However, Desmatolagus would have 
had as good a chance of being preserved as Megalagus or Paleolagus 
if it had been present on the plains during the Chadronian. 

Ischyromys is the same size as Titanotheriomys, and had it been on 
the plains during the Chadronian it would have had as good a chance of 
being preserved as the latter. The occurrence of Ischyromys in the 
Cypress Hills does not necessarily indicate that the fauna was mixed 
(Wood, 1937, p. 193), as Lambe (1908, p. 8) supposed, but could indicate 
temperature zoning on the plains during Lower Oligocene time. The 
large ground squirrels of the genus Citellus (sensu lato) are, with very 
few exceptions, restricted to the mountains and the Boreal Zone of the 
plains. The Cypress Hills are far enough north to be climatically 
equal to Pipestone Springs regardless of what was the Lower Oligocene 
climate as a whole. 

The following genera have been omitted from table 4 because our 
knowledge of them in the Lower Oligocene is inadequate for this type of 
study: Talpa?, Kentrogomphios, Sinclairella, Manitsha, Ardynomys, 
Macrotarsius, Hyaenodon, Pseudopterodon, Aepinacodon, Bathygenys, 
Limnetes, Heteromeryx, Pseudoprotoceras, Hotylopus. 


CANYON FERRY FOSSIL VERTEBRATES—WHITE 435 


In summary, the available data show that there was environmental 
(and, in a sense, climatic) zoning between the mountains and the 
plains in Lower Oligocene times. The faunal differences embrace 
larger systematic categories (Felidae and Entelodontidae) than the 
same areas do today (before civilization changed the picture). Both 
the immigrants and the indigenous faunas exhibit nearly the same 
degree of difference. The only change from Lower to Middle Oligocene 
was that the mountain forms were able to invade the plains and most 
of these were immigrants. None of the plains forms were successful in 
invading the mountains between Lower and Middle Oligocene times. 


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Wuirtt, T. E. 
1947. Additions to the Miocene fauna of North Florida. Bull. Mus. Comp. 
Zool., vol. 99, No. 4, pp. 497-515. 
WIitson, Ropert W. 
1949. On some White River fossil rodents. Carnegie Inst. Washington 
Publ. 584. : 
Woop, A. E. 
1936-1941. See Scott, Jepsen, and Wood. 
Woop, A. E., and Witson, R. W. 
1936. A suggested nomenclature for the cusps of the cheek teeth of rodents. 
Journ. Paleont., vol., 10, No. 5, pp. 388-391. 


U.S. GOVERNMENT PRINTING OFFICE: 954 


INSTITUTION 


SMITHSONIAN 
U. S, NATIONAL MUSEUM 


Vol. 103 Washington: 1954 No. 3327 


A REVIEW OF THE LABRID FISH GENUS WETMORELLA 
WITH DESCRIPTIONS OF NEW FORMS FROM THE TROPI- 
CAL INDO-PACIFIC 


By Leonarp P. Scautrz and N. B. MarsHatu' 


Fowler (Fishes of Oceania, suppl. 1, p. 358, 1931) and Weber and 
de Beaufort (Fishes of the Indo-Australian Archipelago, vol. 8, p. 82, 
1940) list Wetmorella philippina Fowler and Bean (U.S. Nat. Mus. 
Bull. 100, vol. 7, p. 211, pl. 17, 1928, type locality Philippine Islands) 
as a synonym of Cheilinus fasciatus (Bloch). We have studied the 
types of Wetmorella philippina (fig. 1) and conclude without any 
doubt that these specimens are generically distinct from Chewlinus 
and are remarkably distinct from C. fasciatus or any other species of 
labrid currently referred to that genus. The generic characters that 
distinguish Wetmorella from other genera in the Labridae are the 
large scales on the head, which form a pattern not occurring in any 
other genus of labrid fishes, and the acutely triangular head. Fowler 
and Bean (op. cit., pp. 211-212) had six specimens from the Philippines, 
but only four of them, in our opinion, are philippina (holotype: 
USNM 89968 from Little Santa Cruz, Zamboanga; paratypes: 
USNM 93503 from Port Langcan, Palawan, USNM 93505 from 
Atulayan Island, Philippines, and USNM 93528 from Cape Kait, 
Libani Bay, Celebes). The other two specimens represent un- 
described species, and these, along with additional specimens from the 
Marshall Islands and the Red Sea, are described as new. 


1 Curator of Fishes, British Museum (Natural History). 
274881—54 ‘ 439 


440 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Genus Wetmorella Fowler and Bean 


Wetmorella Fowler and Bean, U. S. Nat. Mus. Bull. 100, vol. 7, p. 211, 1928. 
(Genotype, Wetmorella philippina Fowler and Bean.) 

The genus Wetmorella is characterized by dorsal rays [X or X, 
9 or 10; anal III,8; pectoral i1,9 or 10; branched caudal rays 6+5; 
lateral line interrupted with 13 to 15+6 or 7 pores to base of caudal 
fin. Jaws equal or nearly so; premaxillary protractile; teeth short, 
conical, in a single row in both jaws, those near front of both jaws 
becoming gradually enlarged, the two pairs nearest tip of jaws largest; 
gill membranes broadly joined across isthmus, forming a free fold; 
head with a distinctive scale pattern composed of large, characteristi- 
cally shaped scales, arranged in a pattern similar to that shown in 
figure 53. The chief variation in scales on the head is that there may 
be 2 median scales on the snout instead of 1; cheek with 1 or 2 rows 
below which on subopercle may occur another row and a single scale 
below the latter; 2 or 3 rows of scales behind eye, including the gill 
cover; a row of large scales occurs above dorsal lateral line, then a 
second row along spiny dorsal fin, mostly covering the spines except 
tips, becoming much smaller along soft dorsal rays, almost disappear- 
ing on base of last ray; anal fin with a similar sheath of scales; basal 
half of caudal fin enclosed in large scales; axillary pelvic scale present, 
short; interorbital space flattish and a little convex; dorsal profile of 
head nearly straight, forming an angle of 40° to 55° with ventral 
contour of head; maxillary covered by preorbital when mouth is 
closed. A blackish ocellate spot in pelvics and at rear of soft dorsal 
and anal fins; white bar behind eye and on caudal peduncle. 


Key to the species of Wetmorella 


la. Greatest width of white bar between rear of bases of soft dorsal and of soft 
anal fins across caudal peduncle is contained about 2.0 to 4.7 times (3.8 to 
4.7 in Red Sea specimens) in least depth of caudal peduncle; white bar on 
caudal peduncle completely encircles it; young only, with white bar from 
front of spiny dorsal through pectoral base, thence to pelvic base; white 
bar behind eye meets its fellow near occiput. 
2a. Caudal fin plain dusky, except in smallest specimens there occur 2 narrow 
black cross bars, remainder of fin pale or white; some scales on middle 
of sides have black dots; greatest depth about 2.4 to 2.5 in standard 
NON Gt 2228s cee = oe eee a er Wetmorella ocellata, new species 
2b. Caudal fin with a few black spots on middle rays, at about three-fourths 
their length distally; greatest depth 2.4 to 3.2. 
3a. Greatest width of peduncular white bar 2.3 to 3.4 in least depth of 
caudal peduncle; no white bar in front of ocellate spot in soft dorsal 
and in front of soft anal fins (fig. 52). 
Wetmorella philippina phiiippina Fowler 
3b. Greatest width of peduncular white bar 3.8 to 4.7 in least depth of caudal 
peduncle; white bar present in front of ocellate spot in soft dorsal and 
in soft anal fins (pl. 12, fig. A). 
Wetmorella philippina bifasciata, new subspecies 


GENUS WETMORELLA—SCHULTZ AND MARSHALL 44] 


1b. Greatest width of white bar on caudal peduncle between rear of bases of soft 
dorsal and of soft anal fins contained about 6 to 10 times in least depth of 

caudal peduncle; greatest depth about 2.8 to 2.9. 
4a. A white bar from behind ocellate spot in soft dorsal passes in front of 
ocellate spot in soft anal fin; a white bar from bases of third and 
fourth dorsal spines passes behind pectoral base, thence to just 
behind pelvic base; posterior third of caudal fin with a black band, 
but rear margin of caudal fin is edged with white; the white bar on 
caudal peduncle occurs as a saddle ventrally and does not extend 

on dorsal part of caudal peduncle. 

Wetmorella albofasciata, new species 
4b. No white bar passing between ocellate spot in soft dorsal and that in 
soft anal; a narrow, white bar extends from in front of ocellate spot 
in soft dorsal to in front of that in soft anal fin; caudal fin plain 
dusky, edged with white distally; a white bar between orbits in the 
interorbital space_.__---_--- Wetmorella triocellata, new species 


Wetmorella philippina bifasciata, new subspecies 


Figure 53; PLATE 12, Figures A, B 


Holotype—BM 1951.9.18.1, Red Sea, Suakin, Anglo-Egyptian 
Sudan, ‘Manihine’ Collection, January 13, 1951, taken by use of 
derris root (6 percent rotenone) from pieces of coral growing on sea 
wall surrounding Suakin, at depth of 3 feet, standard length 50 mm. 

Paratype-—BM 1951.9.18.2, taken with holotype and bearing same 
locality data, standard length 41.5 mm. 

Deseription.—Certain counts and measurements are recorded for 
the holotype and paratype in tables 1 and 2. 

Body compressed, greatest depth opposite middle of spiny dorsal 
base; snout normal; dorsal profile of head straight or nearly so, 
forming an angle of 52 to 55° with ventral contour of head and body; 


Ficure 52. Holotype of Wetmorella philippina philippina Fowler and Bean (USNM 89968) 
from the Philippines. 


442 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


TABLE 1.—Counts recorded for species of Wetmorella. 


erie Lateral line pores 

Dorsal rays Anal | Pectoral rays) |. 
Species ay Dor- | Ven- 
: sal | tral | Upper | Lower 
lobe | lobe 


W."philippina philippina__.| 3| 1| 1| 3] 4] 4] 7] 1] 6 4 4] 21) Both etal 
philippine bifasciata-.------ Ah Peed eA i ea er) abel ow 2 7 ee ee se Se 45 ee 
Ocei ati Som. os Fee at rot bal seas WP 12h 123 12 12 3 3 |- ta Veto 4 
albofasciata _-_-.------------ 1 ees Pee 1 1 1 ? a aera} 1 pL est fe ae 1 eee 1 
tridcellatas 2 fb ns 1 Tg] ee [Cole 1 1 1 Ayee-Ae 2 1 Piss se 1 ee 


Ficure 53. Sketch of the arrangement of the scales on the head of holotype of Wetmorella 
philippina bifasciata, new species. A, B, holotype; C, paratype. 


interorbital space slightly convex, nostrils small, the anterior one 
tubular; jaws equal or nearly so; scales on head as illustrated in 
figure 53. 

Since the other morphological characteristics of Wetmorella philip- 
pina bifasciata are so similar to W. p. philippina it is not deemed 
necessary to redescribe them here. 

Color in alcohol—Background coloration brownish with a narrowish 
white bar just behind ocellate spot passing through rear base of soft 
dorsal, thence across caudal peduncle through rear of anal base behind 
ocellate spot in soft anal fin; another white bar, much less distinct, 
extends from in front of ocellate spot in soft dorsal fin across body to 
in front of ocellate spot in anal fin; a narrow, white bar extends from 
occiput past rear of eye to lower posterior corner of gill cover; caudal 
fin with about 7 or 8 black spots; pelvics black distally. 

Remarks.—This new subspecies from the Red Sea differs from 
W. p. philippina in having a narrower peduncular white bar and a 
white bar in front of the ocellate spot in dorsal and anal fins. 

Named bifasciata in reference to the two white bands, one in front 
and one behind the ocellate spots. 


GENUS WETMORELLA—SCHULTZ AND MARSHALL 


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444 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Wetmorella ocellata, new species 


PiatTEe 12, Figures D, E 


Holotype—USNM_ 112368, Rongelap Atoll, Kieshiechi Island, 
north end, lagoon coral head, depth 20 feet, July 24, S-46-285,? Brock 
and Herald, standard length 38.6 mm. 

Paratypes—USNM 112372, Bikini Atoll, coral heads in eastern 
end of lagoon, depth 20 to 25 feet, March 26, S-46-42,? Brock and 
Schultz, 2 specimens, 25 mm. and 49 mm.; USNM 112371, Bikini 
Atoll, off Amen Island in lagoon, depth 30 feet, August 4, S-46-307,? 
Herald and Brock, 1 specimen, 50 mm.; USNM 112370, Bikini 
Lagoon, 100 yards off Airy Island, depth 20 to 40 feet, August 7, 
S—46-308,? Brock and Herald, 6 specimens, 23 mm. to 57 mm. Bikini 
Atoll, reef between Amen and Bikini Islands in lagoon, depth 30 feet, 
July 31, 1947, Donaldson and Welander, 1 specimen, 48 mm.; USNM 
112369, taken with holotype and bearing same data, 1 specimen, 41 
mm.; USNM 112373, Bikini Atoll, Amen Island, August 21, 1947, 
Univ. Washington, 1 specimen, 43 mm. 

Description.—Dorsal rays [X,10; anal III,8; pectoral ii,10; pelvics. 
1,5; branched caudal fin rays 6+5; pores in lateral lines 14 or 15+6 
or 7; scales above lateral line 2, below lateral line to anal origin 6; 
vertical scale rows 20 or 21; gillrakers on first arch about 6+9. 
(Certain measurements made on the holotype and two paratypes, 
expressed in thousandths of the standard length, are recorded in 
table 2.) 

Body compressed, the greatest depth opposite middle of spiny 
dorsal base; snout normal; dorsal profile of head straight or nearly so, 
forming an angle of 40° to 46° with ventral contour of head and 
body; interorbital space slightly convex; nostrils small, the anterior 
one tubular; a vertical line through rear nasal opening passes through 
front edge of eye; jaws approximately equal; maxillary reaches to a 
vertical line through front nostril; maxillary covered by preorbital 
when mouth is closed; dentary normal; premaxillary protractile; 
teeth short, conical in a single row in both jaws, those near front of 
both jaws becoming gradually enlarged, the 2 pairs nearest tip of 
jaws largest, and, when mouth is closed, those of lower jaw fitting 


EXPLANATION OF PLATE 12.—A, holotype of Wetmorella philippina bifasciata, new sub- 
species (BM 1951.9.18.1) from Red Sea; B, paratype of same subspecies from same 
locality; C, holotype of W. albofasciata, new species (USNM 93504) from the Philip- 
pines; D, holotype of W. ocellata, new species (USNM 112368) from Rongelap Atoll, 
Marshall Islands; EZ, paratype of W. ocellata, new species (USNM 112372) from Bikini 
Atoll, 25 mm. standard length. 


E 2 Collecting station in Operation Crossroads, 1946 (see U. S. Nat. Mus. Bull. 202, 1958). 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 12 


FoR EXPLANATION SEE FACING PAGE. 


s 


GENUS WETMORELLA—SCHULTZ AND MARSHALL 445 


between the two opposite teeth in upper jaw; gill membranes broadly 
joined across isthmus and forming a free fold; head with a distinctive 
scale pattern composed of large and definite shaped scales as follows: 
Cheek with a single row of enlarged scales below which on subopercle 
is another row, and a single scale below the latter; 3 rows behind eye, 
including gill cover, dorsal surface of head scaled forward to snout 
just in front of orbits, the two anteriormost scales median in position, 
then 3 in middle of interorbital space, followed by a pair of larger ones 
between rear of orbits, then about 5 scales to dorsal fin origin; above 
dorsal lateral line is a row of large scales, then a second row along 
spiny dorsal fin, mostly covering the spines except tips, then the row 
of scales is much smaller along soft rays, almost disappearing on base 
of last ray; anal fin with a similar sheath of scales; basal half of caudal 
fin enclosed in large scales; axillary scale of pelvic short; pectoral fin 
reaches to opposite about seventh scale of lateral line; lateral line 
interrupted, beginning again 2 scale rows below on caudal peduncle; 
fourth pectoral ray usually longest; pelvics reaching or nearly reaching 
anus; caudal fin rounded. 

Color in alcohol.—Background coloration light brownish to brownish, 
with a brown-edged white bar across caudal peduncle just behind rear 
of bases of soft dorsal and soft anal fins, and another brown-edged 
white bar just behind eye from side of head to nape; 3 prominent black 
ocellate spots, one at front of soft dorsal, another at front of soft 
anal, and the largest occupying each pelvic fin and the underlying 
part of the body opposite the pelvic fins; no white bar across inter- 
orbital space; middle of upper lip dark barred. The two smallest 
specimens, 23 mm. and 25 mm., probably represent a juvenile color 
pattern—in addition to the white bar behind the ocellate spots there 
is another white bar in front of them that extends from bases of last 
dorsal spines to bases of anal spines; another white band extends from 
first two dorsal spines just behind pectoral base to pelvics; caudal fin 
white with two narrow cross bars, the distal margin of fin white. 

Ecology.—This interesting new labrid was taken only at depths of 
about 20 to 40 feet in the lagoon among coral heads. It was not seen 
in the intertidal zone of the reefs. 

Remarks.—This new species may be distinguished from species in 
the genus Wetmorella by means of the key. Its closest relative is 
philippina philippina from which it differs in lacking black pigment 
spots in the caudal fin; ocellata has a plain dusky caudal fin in the 
adult, and none of the specimens of ocellata has even a trace of black 
spots in the caudal fin. After studying several hundred species of 
fishes of the tropical Indo-Pacific in numerous families we place a 
great deal of confidence in the color pattern differences such as occur 
in the Chaetodontidae, Labridae, Serranidae, and other families. 


446 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Thus we have decided to recognize two new species and one new 
subspecies. Larger series will make it possible to determine the 
exact status of these new forms. Among our 12 specimens of ocellata 
there is little variation in color pattern except that which occurs 
between small (young) specimens and the large adult specimens. 

Named ocellata in reference to the ocellate spots which help to 
characterize all known species of the genus Wetmorella. 


Wetmorella albofasciata, new species 
Prats 12, E 


Wetmorella philippina (in part) Fowler and Bean, U.S. Nat. Mus. Bull. 100, vol. 
7, p. 212, 1928 (specimen from Mapbul Island, Philippines). 

Holotype—USNM 93504, Mabul Island, Philippines, Albatross, 
September 29, 1909, standard length 36.4 mm. 

Description.—Dorsal rays IX,10; anal JII,8; pectoral 1i1,10-1,10; 
pelvics I,5-I,5; branched caudal 6+5; pores in lateral line 15++-7 with 
2 scales above and 6 below to anal origin; vertical scale rows 21. 

Certain measurements made on the holotype and expressed in 
thousandths of the standard length are recorded in table 2. 

Since the morphological characteristics of albofasciata are so similar 
to those of ocellata it is not deemed necessary to repeat them here 
except to point out that this species is more slender, its greatest depth 
about three times in the standard length. 

Color in alcohol—Background coloration brownish, with a narrow, 
white bar from just behind ocellate spot in soft dorsal, passing across 
body in front of ocellate spot in soft anal fin, ending on front of anal 
fin; peduncular white bar confined to lower half of caudal peduncle, 
not extending much above peduncular lateral line; a short, white bar 
occurs in front of ocellate spot in soft dorsal and ends near dorsal 


Ficure 54. Holotype of Wetmorella triocellata, new species (USNM 93529) from the 
Philippines. Drawn by Mrs. Aime M. Awl. 


GENUS WETMORELLA—SCHULTZ AND MARSHALL 447 


lateral line; another white bar begins near base of third dorsal spine, 
curves ventrally, and passes about 2 scales behind pectoral base, 
thence to behind black area near base of pelvics; distal fourth of cau- 
dal fin blackish edged with white on rear margin and the black caudal 
spot edged with white anteriorly; pelvics blackish except edged with 
white and body next to pelvics blackish. 

Remarks.—This species may be separated from all others referable 
to the genus Wetmorella by the white bar that passes across body 
between the ocellate spots in soft dorsal and soft anal fins. Named 
albofasciata in reference to the characteristic white bars, 


Wetmorella triocellata, new species 
Fiaure 54 


Wetmorella philippina (in part) Fowler and Bean, U. 8S. Nat. Mus. Bull. 100, vol. 
7, p. 212, 1928 (specimen from Rapurapu Island, June 24, 1909). 

Holotype -—-USNM 93529, Rapurapu Island, Philippines, Albatross, 
June 24, 1909, standard length 39.2 mm. 

Desecription.—Dorsal rays IX,10; anal III,8; pectorals 1,10—-1,10; 
pelvics I,5-I,5; branched caudal rays 6-+-5; pores in lateral line 15-++6, 
with 2 scales above and 6 below to anal origin; vertical scale rows 21. 

Certain measurements made on the holotype, and expressed in 
thousandths of the standard length, are recorded in table 2. 

Since the morphological characteristics of triocellata are so similar 
to those of ocellata, it is not deemed necessary to repeat them here, 
except in regard to greatest depth, which is contained about 3 times 
in the standard length; it is notably a more slender fish than ocellata 
and philippina. 

Color in alcohol_—A narrow, white bar occurs behind the two ocellate 
spots in the median fins and another extends from in front of ocellate 
spot in dorsal to in front of that in anal, ending near base of third 
anal spine; background coloration light brown; a brown-edged pale 
bar from behind eyes meets its fellow dorsally on head; probably an- 
other brown-edged pale bar extends across interorbital space although 
this is faded as are all of the pale bars; caudal fin gradually a little 
darker distally, probably narrowly edged with white. We quote the 
following Albatross color note: ‘‘Pale band behind head extends across 
occiput and bounded by brown line in front and behind its en ie 
extent.” 

Remarks.—This new species is recognizable from others referable 
to the genus Wetmorella by its slender form in connection with a dis- 
tinctive color pattern. It is closest to albofasciata in regard to its 
slender form but differs in not having a white bar on the body passing 
between the two ocellate spots in the median fins. 

Named ¢riocellata in reference to the three ocellate spots that char- 
acterize this group of fishes. 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


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SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1954 No. 3328 


REVIEW OF THE ATLANTIC PERIWINKLES, 
NODILITTORINA, ECHININUS, AND TECTARIUS 


By R. Tucker ABBOTT 


Since the 1942 publication in Johnsonia of ‘‘The Genera Tectarius 
and Echininus in the Western Atlantic’? by Clench and Abbott, 
enough additional data have been assembled to warrant a review of 
this group of tropical, littoral snails. In addition to the anatomical, 
distributional, and biological mformation presented here, there is a 
short discussion of the phylogenetic position of the genus Echininus, 
and some changes in nomenclature. 

It had been noted for some time that the operculum of Echininus 
was multispiral, a feature commonly found among members of such 
families as the Trochidae, Potamididae, and Modulidae but unusual 
for the Littorinidae, which are well known for their paucispiral 
opercula. In a paper that has generally been overlooked, Kesteven 
(1903) removed the genus Echininus (Echinella of Kesteven) from its 
customary position in the Littorinidae and placed it in the Modulidae. 
This has necessitated our making a study of the gross anatomy of 
Echininus, Tectarius, and Modulus. This study was made possible 
through the generosity of Mrs. Germaine L. Warmke, who collected 
and airmailed living specimens of Echininus nodulosus Pfeiffer and 
Nodilittorina tuberculata Menke from Puerto Rico. In the process 
of comparing the latter species with Tectarius muricatus Linné, we 
came to the conclusion that Nodilittorina should be considered as a 
full genus closely related to Melarhaphe or Littoraria rather than as a 
subgenus of Tectarius. 

The nomenclatorial adjustments in this paper involve changing the 
name used in Johnsonia, Tectarius tuberculatus Wood, to Nodilittorina 
tuberculata Menke, and a new and presumably valid genotype desig- 
nation for Nodtlittorina. 

278946—54 449 


450 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Family LirToRINIDAE 
Genus Nodilittorina von Martens, 1897 


In Johnsonia, Clench and I had erroneously considered Noduilit- 
torina as a subgenus of Tectarius. From a study of the shell and 
animal characters, the radula, and the type of egg capsules, it appears 
that this group is much more closely allied to the Melarhaphe or 
Iittoraria subgenera of Littorina than to Tectarius. An obvious 
relationship in shell characters is seen between the Indo-Pacific 
Melarhaphe mauritiana Lamarck and such Nodilittorina as miliaris 
Quoy and Gaimard and picta Philippi, all of which have the peculiarly 
flattened, thin inner columella edge and the axial, zigzag color streaks 
on the whorls. The latter two, miliaris and picta, have strong sculp- 
turing which tends towards the production of small nodules which are 
characteristic of Nodtlittorina. 

The floating egg capsule of Nodilittorina tuberculata Menke and 
Littorina (Melarhaphe) ziczac Gmelin (copied in our fig. 55 from 
Marie Lebour, 1945) are similar in that they are drum-shaped and 
with 6 to 7 spiral lines or ridges on the top surface. These spiral 
ridges are absent in the capsules of Tectarius muricatus Linné and 
Inttorina (Melarhaphe) neritoides Linné. The latter species is from 
the eastern Atlantic and is the genotype of Melarhaphe. Should 
it prove to have a simple, single-pronged penis, as does Tectarius, 
it is likely that such species as L. ziczac Gmelin and L. mauritiana 
Lamarck (which have complicated, Nodilttorina-like penes) do not 
belong to Melarhaphe, sensu stricto. 

Three recent papers have contributed to our knowledge of littorinid 
ege capsules (Ostergaard, 1950; Tokioka, 1950; and Tokioka and 
Habe, 1953.) There appear to be three groups of capsules: (1) 
Helmet-shaped—Littorina littorea Linné (Lebour, 1935, p. 375) and 
Littorina pinctado Wood (Ostergaard, 1950, p. 97); (2) simple drum- 
shaped—Littorina neritoides Linné (Lebour, 1935, p. 375) and 
Tectarius muricatus Linné (Lebour, 1945, p. 465); and (8) drum- 
shaped, with ridges on the top surface—Littorina ziczac Gmelin 
(Lebour, 1945, p. 465) and Noditlittorina tuberculata Menke (Lebour, 
1945, p. 465). 

To the latter group, Tokioka and Habe add the egg capsules of 
three possible littorinid snails. ‘These capsules were given the non- 
binomial names of “‘Zittorina-capsula habei, multistriata, and hagruma.” 
The Japanese capsules differ from those in the Western Atlantic 
species in having concentric instead of spiral ridges on the top surface. 
The “hagruma”’ capsule is unique in bearing on its peripheral surface 
a series of 21 gearlike undulations. 


REVIEW OF THE ATLANTIC PERIWINKLES—ABBOTT 451 


The genus Hamus of Klein seems to have been first validated in 1886 
by R. B. Watson (Scaphopoda and Gasteropoda, in Report on the 
Scientific Results of the Voyage of HMS Challenger . . . , vol. 15, 
pt. 42, p. 576). Although he intended it for the group we know as 
Nodilittorina, I prefer to relegate Hamus to the synonymy of the 
trochid Turcica A. Adams, 1854, by here designating Tectaria mon- 
trouziert Fischer, 1878, as the genotype of Hamus Watson, 1886. For 
identity of that trochid species see Hedley (1901, p. 121). 

Wenz’s (1939, p. 523) genotype designation of Nodzlittorina is 
invalid, since Turbo nodulosus Gmelin is not one of the species included 
by von Martens in 1897 either in name or substance and since Turbo 
(not Trochus) nodulosus Gmelin is a turbinid species. Von Marten’s 
Iittorina nodulosa Pfeiffer is an Hchininus. I hereby designate 
Litiorina pyramidalis Quoy and Gaimard as the genotype of Nodiht- 
torina von Martens, 1897. This species is Trochus nodulosus Gmelin, 
non Solander, 1766. 

Habe (1951, p. 90) followed the error of Clench and Abbott (1942) 
in considering Turbo tuberculata Wood the same as Gmelin’s nodulosus. 
Actually it is the West Indian Echininus, as seen by our revised 
synonymy. Jattorina tuberculata Wood is not included by name in 
von Martens’ genus and cannot be designated as the genotype. 

Below, I have included a revised synonymy of the Western Atlantic 
Nodilittorina tuberculata Menke which Clench and I had erroneously 
listed in Johnsonia as Tectarius tuberculatus Wood. Our error arose 
in not recognizing Wood’s figure as representing the Atlantic Hchininus. 
Although the columella region of the shell illustrated by Wood is very 
close to that found in some Nodilittorina, the nature and number of 
nodules clearly identifies it as Echininus nodulosus Pfeiffer. 


Nodilittorina tuberculata Menke 


Trochus nodulosus Gmelin (non Solander, 1766), Caroli a Linné Systema 
naturae ..., ed. 13, p. 3582, No. 98, 1791 (in part as variety minor). 
(Mari, Americam meridonalem.) 

Litorina tuberculatus Menke, Synopsis methodica Molluscorum ..., p. 25 
(Pyrmonti), 1828 (refers to Gmelin’s minor). 

Intorina thiarella Anton, Verzeichniss der Conchylien. . . , p. 53, 1839. 

Littorina nodulosa d’Orbigny, Mollusques, vol. 1, p. 205 (exclusive of synonymy), 
pl. 14, figs. 11-14, in de la Sagra, Histoire physique, politique et naturelle de 
l’Ile de Cuba, 1841. (Cuba, Martinique, and St. Lucia.) 

Littorina dilatata d’Orbigny, Mollusques, vol. 1, p. 207, pl. 14, figs. 20-23, zn de 
la Sagra, Histoire. . . naturelle. . . de Cuba, 1841. (Havana.) 


Shell—It is easy to confuse this species with Echininus nodulosus 
Pfeiffer, especially since their ranges overlap and their habitats are 
almost identical. N. tuberculata is usually 4 mm. to 5 mm. smaller 
than EH. nodulosus from the same small area. The nodules in each 


452 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


spiral row on the periphery of the whorl are always lined up under 
one another in N. tuberculata, while in E. nodulosus they are not, 
since the upper row bears fewer (and larger) nodules than the lower 
row. 

Sexual dimorphism in the form of shell length was very slight in 
103 specimens examined from Rincon Lighthouse, Puerto Rico. Of 
that number, 38 percent were males; their mean shell length was 7.7 
mm. The mean length of the females was 9.1mm. In Echininus 
the difference in the sizes of the shells of the two sexes was more 
pronounced, but this is a feature which is variable from colony to 
colony in the same species (see Abbott, 1949). 

Animal.—Live specimens from Rincon Lighthouse, western Puerto 
Rico, were examined. The animal is typically littorinid. The 
tentacles are translucent yellowish with a small, circular band or 
ring of black around the region near the distal end. The underside 
of the foot is yellowish to yellowish gray with a distinct, longitudinal, 
indented line dividing the foot into two lateral areas. The furrow is 
more prominent near the center, and disappears towards the posterior 
and anterior ends of the sole. The waves of progression are retrograde 
and ditaxic (see Vlés, 1907, and remarks under Tectarius muricatus). 
The foot progresses somewhat in the manner of a person slowly shuf- 
fling forward in a potato sack. 

The penis is large and located on the right side of the body under 
the base of the right tentacle. On the anterior edge of the penis 
there is a large, bean-shaped, snow-white gland, and just distal to it 
is a short, fairly large, clear accessory flagellum. The main prong 
of the penis is slender and tapering to a point, with a nearly closed 
seminal groove along the posterior edge. The penis of N. tuberculata 
is very similar to that of preserved specimens I have examined in 
Inttorina (Melarhaphe) ziczac Gmelin from the Bahamas. 

Radula.—The ribbon is very long with the unused part coiled up 
like a watch spring in a pocket in the dorsal region behind the head. 
One specimen from Puerto Rico had about 1,125 transverse rows. 
The ribbon is delicate and half as wide as that found in Echininus. 
The central tooth is narrow, with the appearance of having been 
laterally compressed. It bears a large central cusp, and, crowded 
over this, are the two lateral cusps. The lateral and inner marginal 
teeth are massive, each with a large inner cusp and a much smaller 
outer cusp. The outer marginal tooth is smaller, fairly weak; it 
bears 5 denticles in Puerto Rico specimens and 8 denticles in Habana, 
Cuba, specimens. 

Parasitology.—In living specimens examined from the rocky shore 
near Habana, Cuba, several specimens of N. tuberculata were found to 
be heavily infected with single-tailed, two-eye-spotted cercariae whose 


REVIEW OF THE ATLANTIC PERIWINKLES—ABBOTT 453 


aunt] 


Ficure 55.—a, Shell of Echininus nodulosus Pfeiffer (X 1%). b, Shell of Nodilittorina 
tuberculata Menke (X 1%). c, Underside of foot of E. nodulosus (X 2). d, Underside of 
foot of N. tuberculata (X 2). e, Operculum of £. nodulosus (X 2). f, Operculum of N. 
tuberculata (X 2). g, Fecal pellets of £. nodulosus (X 4). h, Fecal pellets of N. tubercu- 
lata(X 4). i, Fecal pellets of Modulus modulus Linné (X 4). 7, Penis of Littorina ziczae 
Gmelin (X 6). &, Floating egg capsule of L. ziczac (diam. 0.20 mm.). /, Egg capsule of 
N. tuberculata (diam. 0.24mm.). m, Egg capsule of Tectarius muricatus Linné (diam. 0.24 
to0.32. After Lebour, 1945). n, Penis of N. tuberculata (X 6). 0, Penis of T. muricatus 
(X 6). 9,49, Penis of E. nodulosus (X 6). 


454 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


identity was unknown to me. They were located in the upper part 
of the whorls in the region of the digestive glands. They may possibly 
be parasites of sea birds. None of the Echininus nodulosus Pfeiffer 
collected in the same area was infected. 

Geographical distribution.—It is interesting that the geographical 
range of this species is identical with that of Yectarius muricatus. 
Both species are coastal rock dwellers, the former living in the im- 
mediate vicinity of tide and splash pools, the latter living anywhere 
from the spray zone, where the rocks are wet only during windy 
weather at high tide, to a point 70 feet from the sea, where the rocks 
are dry except during rainy or stormy periods (Clench and Abbott, 
1942, pp. 2, 3). The Nodzilittorina distribution in the Western At- 
lantic (see fig. 56, a) is Caribbean in its limits with two minor northern 
extensions, one along the southeast coast of Florida, the other in 
Bermuda. There have been no records in the Gulf of Mexico or 
along the eastern coast of South America south of Trinidad, as is the 
case, for instance, with Modulus modulus Linné. 

On the other hand, the geographical range of Echininus nodulosus 
Pfeiffer is much more limited (fig. 56, 6) and consists of a compact 
oval area which includes southeast Florida, the Bahamas, and the 
Greater Antilles. Despite numerous museum records for other lit- 
toral species, the locally common Hchininus has not been recorded 
from Bermuda, the Lesser Antilles, or the Caribbean shores of Central 
or South America. The reasons for this difference in distribution is, 
as yet, unexplained, but the geological history and the life history of 
these two groups, when known, will likely offer a solution. The rela- 
tively dry habitat of Echininus and Tectarius muricatus in contrast 
to the lower and wetter station of N. tuberculata precludes any corre- 
lation between the immediate, ecological niche and the distributional 
range. However, food habits, dispersal factors by other animals, 
and enemies are unknown. Both Nodilittorina and Tectarius have 
been found to expel single, floating egg capsules (Lebour, 1945), but 
the manner of egg deposition is unknown for Echininus. 

The facts published by Mattox (1949) concerning the ecological 
station (bio-stratification) and resistance to drying conditions (ex- 
siccation) of Nodilittorina tuberculata (as Tectarius tuberculatus) in 
Puerto Rico are open to question since they may be based upon ob- 
servations on two genera. LEchininus is commonly found in company 
with Nodilittorina along the rocky shore of the western end of Puerto 
Rico. The two are easily confused and the author makes no mention 
of the common Echininus in his report on the eight littoral species of 
that area. 

New records.—Bauamas: Andros Island; Aklin Island; New 
Providence Island; Great and Little Inagua Islands; Mariguana 


REVIEW OF THE ATLANTIC PERIWINKLES—ABBOTT 455 


Zan 
(%) 
PA 
a 


—~Y NODILITTORINA 


Ficure 56.—Distribution in the West Indies of Nodilittorina tuberculata Menke, Echininus 
nodulosus Pfeiffer and Tectarius muricatus Linné (see locality records in text). 


456 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 108 


Island; East and Grand Caicos Islands; Cotton Cay, Turks Islands 
(all P. Bartsch). CusBa: Cape Cajon; Cabafias; Rio Ojo de Toro, 
Oriente. Hispanioza: Torbeck, Dept. du Sud, and Saltrou, Dept. 
de l’Ouest, Haiti. Jamaica: Little River, Trelawney; Dry Harbor 
Cave, St. Ann; Great Pedro Bay, St. Elizabeth; Ora Cabesa, St. 
Mary; Hector’s River, Portland; Little Cayman Island (all C. R. 
Orcutt). Purrro Rico: Rincon Lighthouse, Mona Passage (G. L. 
Warmke and N. T. Mattox); Aguadilla, Porto Real; Mona Island. 
Lesser Antities: Villa, St. Vincent; Marigot, Dominica; Guade- 
loupe. CrnrraL America: Fort Sherman, Col6én, Panama (L. D. 
Sayers). SourH America: Santa Marta, Colombia. (All records 
USNM.) 

I am also adding here a revised synonymy of the Indo-Pacific 
sibling species, NV. pyramidalis Quoy and Gainard. I have compared 
material from northern Australia (the region of the type locality) 
with specimens from the East Indies, the Philippines, and the Ryukyu 
Islands and do not believe, on the basis of the material at hand, that 
N. vilis or N. malaccana should be recognized even as subspecies. I 
have not included subnodosa Philippi, 1847, in the synonymy for lack 
of sufficient number of specimens from its Red Sea type locality. It 
may well be a good subspecies. The Formosan specimen figured by 
Habe (1951, pl. 14, fig. 1) is probably not the Red Sea species, despite 
its close resemblance. 


Nodilittorina pyramidalis Quoy and Gaimard 


Trochus nodulosus Gmelin (non Solander, 1766), Caroli a Linné Systema naturae 
. , ed. 18, p. 3582, No. 98, 1791 (refers to Martini and Chemnitz, Neues 
Systematisches conchylien-cabinet, vol. 5, pl. 168, figs. 1545, 1546 (of 
specimens from Cook’s voyages ‘“‘aus den Sudlandern.’’) (“Oceano australi.”’) 
Turbo trochiformis Dillwyn (non Brocchi, 1814), A descriptive catalogue of re- 
cent shells . .. , vol. 2, p. 826, 1817 (refers to Gmelin, 1791, p. 3582, and 
Martini and Chemnitz (loc. cit.). (‘‘SSouthern Ocean.’’) 
Littorina pyramidalis Quoy and Gaimard, Zoologie, vol. 2, p. 482, pl. 33, figs. 


12-15, in d’Urville, Voyage de .. . l’Astrolabe . . . pendant... , 1833. 
(Jervis Bay, Australia.) 

Iitorina vilis ‘“Menke”’ Philippi, Abbildungen und Beschreibungen .. . Con- 
chylien ... , vol. 2, p. 145, pl. 2, fig. 21, 1846. 

Litorina malaccana Philippi, Abbildungen und Beschreibungen . . . Conchylien 


... , Vol. 8, p. 51, pl. 6, fig. 17, 1847. (Pulo Pinang.) 
Litorina cecillei Philippi, Zeitschr. Malakoz., vol. 8, No. 2, p. 78, 1851. (Liew- 
kiew = Ryukyu Islands.) 
Type locality —Jervis Bay, Australia. 
Range.—India and Ceylon, Siam, the Philippines, and East Indies, 
south to New South Wales, Australia, and north to Honshu, Japan. 


REVIEW OF THE ATLANTIC PERIWINKLES—ABBOTT 


sung 
Ll 
it 


Ficure 57.—Radulae of, a, Echininus nodulosus Pfeiffer, b, Nodilittorina tuberculata Menke, 
c, Modulus modulus Linné, and, d, the central tooth of Tectarius muricatus Linné. (Arrows 
point to side views of teeth.) 


458 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Genus Echininus Clench and Abbott, 1942 


Echininus, sensu stricto, is limited to the Indo-Pacific. It differs 
from the subgenus Tectininus Clench and Abbott in having an um- 
bilicus and in having a more spinulose sculpturing. It was formerly 
known as Nina Gray, 1850 (non Horsfield 1829, non Nina Gray 1855). 
Echinelia of Kesteven, 1903, and some other authors is this genus, 
but Echinella Swainson is a synonym of Tectarius Valenciennes. 
Genotype, by original designation, is Trochus cumingi Philippi. 


Subgenus Tectininus Clench and Abbott, 1942 


Many of the subgeneric characters listed here may well be those 
of the genus as a whole, but we have not had an opportunity, as yet, 
to examine the genotype of Echininus, sensu stricto, in detail. The 
genotype, by original designation, is Hchininus nodulosus Pfeiffer. 

The general anatomy, both internal and external, is typically lit- 
torinid. Operculum loosely multispiral with about 6 whorls (trochid 
opercula of the same diameter, such as in Calliostoma and Margarites, 
have 11 to 13 whorls). Shell whorls roughly bicarinate, with the 
upper carina bearing about one-third fewer nodules than the lower 
carina. Columella short, and not shelved or protruding downward 
beyond the base of the outer lip as in Nodilittorina. Radula ribbon 
with about 800 to 1,250 transverse rows of teeth. Unused ribbon 
coiled in about 16 close turns and located in the dorsal region behind 
the head. Central tooth extremely small and slender. Lateral teeth 
very large, strong, and with a strong, hooked, single cusp. Inner 
marginal tooth with a single, large cusp. Outer marginal tooth half 
the length of the lateral one, and tri-denticulate. Foot weakly di- 
taxic, with retrograde waves. Penis large and located on the right 
side of the head just below and slightly posterior to the base of the 
right tentacle. Penis tri-lobed and with a seminal groove running 
along the posterior edge. In the area where the accessory prong arises 
there are numerous, small papillae on the penis. In the female, the 
ovaries are widespread through the digestive gland and are grass- 
green in color. The area of the prostate gland in the males is bright 
Prussian blue. 

Echininus nodulosus Pfeiffer 
Turbo tuberculatus Wood (non Pennant, 1777), Index testaceologicus, suppl., 
p. 19, pl. 6, fig. 30, 1828. 


Litorina nodulosa Pfeiffer, Arch. Naturg., vol. 1, p. 357 (exclusive of synonymy), 
1839. (Cuba.) 


Litorina scabra Anton, Verzeichniss der Conchylien .. . , p. 53, 1839. 
Ittorina antoni Philippi (new name for Littorina scabra Anton, non L. scabra 
Linné, 1758, of authors), Abbildungen und Beschreibungen . . . Con- 


chylien . .. , vol. 2, p. 145, pl. 2, fig. 18, 1847. 


REVIEW OF THE ATLANTIC PERIWINKLES—ABBOTT 459 


Litorina (Tectarius) pfeifferianus Weinkauff (new name for Littorina nodulosa 
Pfeiffer, non Littorina nodulosa Gmelin of authors), Litorina, in Martini 
and Chemnitz, Systematisches Conchylien-Cabinet, vol. 2, pt. 9, p. 46, pl. 5, 
figs. 15, 16, 1882. 


Shell—Perhaps the most distinctive feature of the shell of this 
species is the nature of the rows of nodules. As in Nodalittorina 
tuberculata, the periphery of the last whorl bears two prominent rows 
of large nodules, thus giving the shell a slightly bicarinate appearance. 
There are five or six minor rows of smaller beads. But especially 
notable in Echininus are the fewer and larger nodules in the upper 
major row. A count in 30 specimens revealed that, on the average, 
there are 3 or 4 fewer nodules in the upper row. No significant differ- 
ence in this feature could be found between the males and the females. 
(In 15 males, the upper row, on the average, bore 11.3 nodules with a 
range of 9 to 14; the lower row had 14.9 nodules with a range of 11 to 
17. In 15 females, the upper row bore 10.9 nodules with a range of 
9 to 13; the lower row bore 13.0 nodules with a range of 10 to 15.) 

Measurements of shell length were compared between 18 adult 
males and 21 females, and it was found that there is a slight sexual 
dimorphism in which the mean length of males is 13 mm. and that of 
the females 15 mm. The overlap in shell size, however, is too great 
to permit distinction of sex on this character alone. 

I notice that the shells of some of the females are more eroded than 
those of the males, and in this connection there is need for further 
study on the wanderings of the females and their possible subjection 
to conditions slightly different from the males. M. Lebour (1945) 
believes that the amphibious Littorinids of Bermuda seasonally 
descend from their positions high in trees or from rock cliffs to the 
edge of the ocean to deposit their eggs. Whether the males also 
migrate is unknown. 

Animal.—Our studies were made on living specimens from Rincon 
Lighthouse, western Puerto Rico. The animal is typically littorinid, 
as discussed under the remarks concerning the subgenus. The mantle 
edge is smooth, slightly swollen, and yellowish; the remainder is clear. 
The underside of the foot is yellowish gray, but not as yellow as in 
Nodilittorina. The longitudinal fissure on the sole of the foot is very 
weak and limited to the posterior half. The retrograde (front to 
back) waves at the anterior third of the sole are usually several and 
somewhat confused, but as they proceed posterior and reach the 
region of the weak, central fissure, they become stronger and take on 
a ditaxic, lateral division. 

The penis is large, prominent, and located on the right side of the 
body of the males. The accessory flagellum is cylindrical, larger than 
that found in Nodilittorina, and located one-third from the distal end 


460 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


of the penis. On both sides of the penis, in the region near the acces- 
sory flagellum, the surface bears numerous, small, fleshy, opaque-white 
papillae. These are absent in Nodilittorina and Tectarius muricatus. 
The posterior edge of the penis bears a narrow, nearly sealed 
seminal groove which ends near the distal end of the penis, and which 
has its origin on the side of the body some 2 mm. from the base of the 
penis. The region of the prostate gland on the right border of the 
mantle is cobalt blue in color. In mature females, this region bears 
the swollen, elongate, opaque-white accessory gland. 

Radula.—The radula of Echininus nodulosus Pfeiffer (fig. 57, a) is 
very distinctive, although it conforms in general pattern to those 
found in the family Littormidae. The reduction of the central tooth 
is most remarkable in that it has become a short, thin, sliverlike 
tooth. The main function of rasping has been taken over by the 
greatly enlarged, coarse lateral teeth. The inner and especially the 
outer marginal teeth are somewhat reduced. The tendency to reduce 
the central tooth appears to be a development arising perhaps through 
the Nodihttorina stock. The radula of Echininus cumingi from the 
Indo-Pacific has a reduced central tooth (fide Troschel, 1858, pl. 11, 
fig. 7) and enlarged lateral teeth. Among the Nodilittorina, our 
Atlantic N. tuberculata Menke has a strongly compressed, elongate 
central tooth (fig. 57, 6). The Indo-Pacific N. pyramidalis has a 
much less reduced central tooth which closely resembles that in the 
members of the subgenus Melarhaphe. It may be noted that mem- 
bers of such species as Littorina (Littorina) littorea Linné and irrorata 
Say have a well-developed, almost square central tooth. 

The radula ribbon of Echininus nodulosus measured 115 mm. in 
length in one specimen, 110 mm. in another. This is 7 to 8 times the 
length of the shell. The ribbon is twice the width of that found in a 
Nodtlittorina tuberculata Menke of the same shell size. The number 
of transverse rows is about 1,250. In N. tuberculata there were about 
1,225 rows, in Tectarius muricatus 1,500 rows in a ribbon 67 mm. in 
length. Pelseneer (Mollusca, Treatise in Zoology, 1906) reports 
3,600 rows in Jattorina littorea. I do not know if there is any phylo- 
genetic or ecological significance to these various number of rows. 

Geographical Distribution.—The range of this species is presented in 
figure 56, 6, and its restricted distribution is discussed in the remarks 
under Nodilittorina tuberculata. The species is usually abundant 
where it occurs. 

New records.—F.oripa: Stock Island, Key West (C. I. Aslakson) ; 
Indian Key; Fortune Island (P. Bartsch). Banamas: Nassau, New 
Providence Island (H. Dodge); South Bight, Andros Island (P. 
Bartsch); Pimlico Island, Racoon Cay, Ragged Islands (P. Bartsch). 
Cupsa: Cape Cajon; Santa Cruz; Cienfuegos Harbor; Santiago de 


REVIEW OF THE ATLANTIC PERIWINKLES—ABBOTT 461 


Cuba. Jamatca: Robins Bay, St. Marys; Montego Bay, St. James; 
Port Henderson; Annotta Bay (all C. R. Orcutt). Huispanroza: 
Morne Rouge, southwestern Haiti; Beata Island (A. Wetmore). 
Puerto Rico: Rincon Lighthouse, Mona Passage (G. L. Warmke 
and N. T. Mattox). Virern Istanps: St. Croix (H. F. Dunn). 
CartBBEAN Istanps: Navassa Island. (All records USNM.) 

Phylogenetic position of Echininus——From the anatomical evidence 
presented here, it appears that there is little doubt that Hehininus has 
many characters that are typically littormid. This is especially true 
in the case of the long radula ribbon, the form of the teeth, the pres- 
ence of an open seminal groove (instead of a closed, internal seminal 
canal) on the penis, and the simple tentacles with the eye set near 
the base. We reject Kesteven’s (1903) placement of this genus in 
the family Modulidae. He did so on two characters—the tooth on 
the base of the columella, and the multispiral operculum. However, 
Kesteven was in error in stating that members of the genus Echininus 
(Echinella, as he called it) have a small tooth at the base of the col- 
umella. It is only in the Tectarius, sensu stricto, group (with 
paucispiral opercula) such as 7. coronarius Lamarck, that we find a 
basal tooth. This leaves only the multispiral operculum as a character 
in common with Modulus, and although this character may be of 
convenient generic value in some cases, it certainly is not always 
a family character. 

I believe that the following characters found in Modulus, and 
absent in Echininus, exclude the latter from the family Modulidae: 
Eyes located half way up the length of the tentacles, small digitations 
along the border of the mantle edge (Abbott, 1944, pl. 1); female with 
an “ovipositor” organ on the right side of the body; radula relatively 
short (100 to 150 transverse rows) with a thin, oval, 7-denticled 
central tooth; a lateral and two marginal teeth which are denticulated 
(the entire radula closely resembles that in the Rissoacea). AlI- 
though not necessarily of phylogenetic importance, it may be pointed 
out that the feces of many, if not all, Littorinidae are relatively short 
(2 or 3 times as long as wide) and are lined up in the rectum, one 
directly behind the other, while in Modulus modulus Linné the feces 
are quite long (5 or 6 times as long as wide) and are closely packed 
side by side at an oblique angle in the rectum, as in the Thiaridae. 

In summary of the position of Echininus, I am inclined to consider 
it a specialization of the ancestral stock of the Melarhaphe group in 
the family Littorinidae which has shown a tendency towards the 
abortion of the central tooth in the radula, an enlargement and closer 
juxtaposition of the marginals, the reduction of the ditaxic fissure in 
the foot, a reduction in the size of the osphradium, and the increase 


462 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


in the number of whorls in the operculum to a degree where it has 
become multispiral. 

It should be pointed out that Kesteven included the genus Peasiella 
Neville in the family Modulidae. Although further anatomical study 
is needed on this genus, I would be inclined (with the radular and 
opercular characters presented by Kesteven, 1903, p. 633) to include 
Peasiella in the family Littorinidae and in a higher phylogenetic 
position than the genus Echininus. Kesteven also erected a new 
family, the Risellidae, for the inclusion of Risella Gray (now Bembicium 
Philippi) and Aisellopsis Kesteven, 1902 (not Cossmann, 1908, which 
was renamed Risellordea Cossmann, 1909). The new family name, 
Bembiciidae, was introduced by A. W. B. Powell (1937, p. 67). 
Before accepting this family, it would be best to have an anatomical 
study made, and, in the meanwhile, allow it to remain in the family 
Littorinidae as do J. Thiele and W. Wenz. 


Genus Tectarius Valenciennes, 1833 
Subgenus Cenchrites von Martens, 1900 


Tectarius muricatus Linné 


Tectarius muricatus Linné, Systema naturae, ed. 10, vol. 1, 1758. 

Observations on living specimens from Vedado, Habana, Cuba, 
show that the animal is typically littorinid. The tentacles are one- 
third longer than the extended proboscis, are cylindrical and tapering, 
translucent gray in color with faint circular bars of brown. The 
proboscis is dark brown. The mantle edge is thickened, cream yellow 
in color, and slightly undulatory. The sides of the foot are light 
slate-gray with numerous, fine, clusters of chalk-white, embedded 
granules. Underside of foot is slate gray. 

The mode of foot progression is ditaxic with a few simultaneous 
retrograde waves; that is, the sole of the foot is divided down its 
length from anterior to posterior by a faint fissure, and each side 
half acts independently of the other much in the manner of a person 
shuffling forward with his feet in a potato sack. On each half, a 
dark wave appears at the anterior end at the same time and moves 
towards the posterior end (retrograde wave). A bibliography con- 
cerning this subject is given by H. W. Lissmann (1945). This same 
type of progression was reported for Litiorina littorea Linné (Vlés, 
1907; Parker, 1911), Littorina sazatilis Olivi (as L. rudis Maton), and 
Nodilittorina tuberculata Menke (as Tectarius nodulosus from Bermuda) 
(Parker, 1911). 

The penis of Tectarius muricatus is the simplest of the littorinids I 
have examined from the Western Atlantic, and, except for the absence 
of sawlike frills on one edge, it is not unlike that of Littorina littorea 


REVIEW OF THE ATLANTIC PERIWINKLES—ABBOTT 463 


Linné. It is located at the usual position on the right side of the 
body, and consists of a simple bent, tapering prong which bears a 
thin, seminal groove along its posterior edge. 

The radula in one specimen was 67 mm. in length, with about 1,500 
transverse rows of teeth. As in Littorina, the unused rows are coiled 
tightly like a watch spring in a pocket on the dorsal region behind 
the head. The central tooth is about half as wide as long, not unlike 
that in Nodilitiorina, and bears a large central cusp with a smaller 
cusp on each side. (See fig. 57, d.) 

The shell is adequately illustrated in Johnsonia by Clench and 
Abbott (1942). 

Geographical distribution.—In addition to the records noted on the 
accompanying map (fig. 56, c) of the West Indian region, I am listing 
a few new ones which represent either extensions of the range, addi- 
tional major islands, or the filling in of large gaps (for other records, 
see Clench and Abbott, 1942). Distribution in the Caribbean region, 
lower Florida, and Bermuda is almost identical with that of 
Nodilittorina. 

New records.—Bauamas: South Bight, Andros Island; Little San 
Salvador Island. Cusa: Matanzas; Cardenas Bay; Port Gibara; 
Santiago de Cuba. Hispanioza: Fort Liberte, Dept. du Nord; Cap 
Haitien, Dept. du Nord; Torbeck and Aquin, Dept. du Sud; Santa 
Barbara de Samana. Jamaica: Buff Bay, Portland; Stony Cave, 
St. Mary; Great Pedro Bay, St. Elizabeth; Runaway Bay, St. Ann; 
Montego Bay, St. James; Morant Bay, St. Thomas; Little Cayman 
Brac (C. R. Orcutt). Pusrtro Rico: Mayagitiez; Hamacoa. LxrsszRr 
Antittes: Marigot, Dominica. CARIBBEAN IsLANDS: Curagao; 
Aruba. Crnrrat America: Utilla Island and Roatan Island, 
Honduras; Colén, Panamé; Cartagena and Porto Colombia, Colombia; 
Tucacas, Falcon, Venezuela. (All records USNM.) 


464 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


References 
Assortr, R. Tucker. 
1944. The genus Modulus in the Western Atlantic. Johnsonia (Harvard 
Univ.), vol. 1, No. 14, 6 pp., 2 pls. 
1949. Sexual dimorphism in Indo-Pacific Strombus. Nautilus, vol. 63, 
No. 2, pp. 58-61, 1 fig. 
BEQUAERT, JOSEPH C., 
1943. The genus Littorina in the Western Atlantic. Johnsonia (Harvard 
Univ.), vol. 1, No. 7, 27 pp., 7 pls. 
CLENcH, WiturAMm J., and Asport, R. Tucker. 
1942. The genera Tectarius and Echininus in the Western Atlantic. John- 
sonia (Harvard Univ.), vol. 1, No. 4, 4 pp., 3 pls. 
Hasse, TADASHIGE. 
1951. Littorinidae in Japan. ITlus. Cat. Japanese Shells, vol. 1, No. 4, 
pp. 87-93, 1 pl. 
HEDLEY, CHARLES. 
1901. A revision of the types of the marine shells of the Chevert Expedition. 
Records Australian Mus., vol. 4, p. 121. 
KestEven, H. LercuHron. 
1903. Notes on Prosobranchiata, No. 2—Littorinacea. Proc. Linn. Soc. 
New South Wales, 1902, pt. 4, pp. 620-636, 1 pl. 
Lesour, Marte V. 
1945. The eggs and larvae of some prosobranchs from Bermuda. Proc. 
Zool. Soc. London, vol. 114, pt. 4, pp. 462-489, figs. 
LIssMANN, H. W. 
1945. The mechanism of locomotion in gastropod molluscs. Journ. 
Exper. Biol., vol. 21, No. 1, pp. 58-69, 11 figs. 
Marrox, Norman T. 
1949. Effects of drying on certain marine shells from Puerto Rico. Ecology, 
vol. 30, No. 2, pp. 242-244, 2 figs. 
OsTERGAARD, JENS MATHIAS 
1950. Spawning and development of some Hawaiian marine gastropods. 
Pacific Sci. (Honolulu), vol. 4, No. 2, pp. 75-115, 42 figs. 
PARKER, GrorGeE H. 
1911. The mechanism of locomotion in gastropods. Journ. Morph., vol. 
22, pp. 155-170. 
Powe, A. W. B. 
1937. The shellfish of New Zealand. 100 pp., 18 pls. 
Toxioka, T. 
1950. Droplets from the plankton net: V, New names for egg capsules of 
littorinid gastropods. Publ. Seto Marine Biol. Lab., vol. 1, No. 3, 
pp. 151-152, fig. 6. 
ToxioKka, T., and Hass, TADASHIGE 
1953. Droplets from the plankton net: XI, A new type of Littorina-capsula. 
Publ. Seto Marine Biol. Lab., vol. 3, No. 1, pp. 55-56, fig. 11. 
Viks, F. 
1907. Sur les ondes pédienses des mollusques reptateurs. Compt. Rend. 
Acad. Sci. Paris, vol. 145, pp. 276-278. 
Wenz, WILHELM. 
1939. Gastropoda. Handb. Paliozool., vol. 6, pt. 4, pp. 481-720. [See 
p. 523.] 


U. S. GOVERNMENT PRINTING OFFICE: 1954 


PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


Coa 
fog 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1954 No. 3329 


MAMMALS OF NORTHERN COLOMBIA, PRELIMINARY 
REPORT NO. 7: TAPIRS (GENUS TAPIRUS), WITH A 
SYSTEMATIC REVIEW OF AMERICAN SPECIES! 


By Pattie HERSHKOVITZ 


All known New World species of tapirs occur in Colombia. Repre- 
sentatives of the Brazilian tapir (Tapirus terrestris) were secured in 
northern Colombia by the writer during his 1941-1943 tenure of the 
Walter Rathbone Bacon Travelling Scholarship. The mountain, or 
woolly, tapir (Tapirus pinchaque), originally described from the 
Bogoté region in Colombia, is found in parts of the temperate zones of 
Colombia and Ecuador and does not range into coastal provinces. 
Baird’s tapir (Zapirus bairdii), the largest indigenous land mammal 
of the Neotropical region, is here recorded for the first time from 
South America. The author discovered the coexistence of this species 
with the Brazilian tapir in northwestern Colombia while conducting 
the Chicago Natural History Museum—Colombian Zoological Expedi- 
tion (1949-1952). A woolly tapir was also taken, and it is described in 
this paper. A fourth species, Tapirus indicus, the only other living 
form of tapir, is Asiatic in distribution. 


1 Previous reports in this series have been published in the Proceedings of the U. 8. National Museum as 

follows: 
1. Squirrels, vol. 97, August 25, 1947 

. Spiny rats, vol. 97, January 6, 1948 
. Water rats, vol. 98, June 30, 1948 
Monkeys, vol. 98, May 10, 1949 
. Bats, vol. 99, May 10, 1949 
Rabbits, vol. 100, May 26, 1950 

Egrata;: In No. 5: Bats, measurements given for Glossophaga soricine soricina on page 438 are misleading 
because of a transposition. On line 25, for “‘head and body” read “‘total length,”’ for “‘tail” read ‘‘head and 
body,” for “hind foot” read ‘‘tail,’”’ for “‘ear’’ read “hind foot’’, and add “ear, 15-15 mm.” 


279127—54 1 465 


A ie wb 


ad 


466 PROCEEDINGS OF THE NATIONAL MUSEUM VOU. 103 


Nomenclature 


The generic name Japirus employed here is from Brinnich, 1771. 
Scopoli’s use of Tapirus in 1777 is next available. For rejection of 
Brisson’s Tapirus and other Latin names in his “Regnum Animale,” 
1762, see Hopwood (Proc. Zool. Soc. London, vol. 117, pp. 534-536, 
1947). Tapirus Briinnich is adopted here on the same authority, a 
copy of the ‘‘Zoologiae Fundamenta . .. ,”’ where the generic name 
appears, not being available in this country. Merriam (Science, new 
ser., vol. 1, p. 376, 1895) employed an ingenious device in his attempt 
to validate Tapirus Brisson. He combined his own with Brisson’s 
(Regnum Animale, p. 81, 1762) monomial specific Latin designation 
for ‘Le Tapir” to produce the custom-made binomial Tapirus tapirus. 
This combination is valid, to be sure, but dates from its originator, 
Merriam, 1895, and not from Brisson. The question that has arisen 
over the basic date of publication of the ‘‘Regnum Animale’”’ is en- 
tirely subordinate to the fact that the system of classification em- 
ployed therein is incontrovertibly non-Linnaean. Hence, Brisson’s 
Latin names, really classical rather than technical, are not available. 

Bibliographic references and citations to generic synonyms are given 
under the subgeneric headings. Synonymies under specific headings 
include references to all original descriptions and to selected taxonomic 
works. 


Classification 


Cranial and external differences between living species of American 
tapirs are such as to warrant full generic rank for each of the recog- 
nized forms. Simpson (Bull. Amer. Mus. Nat. Hist., vol. 86, pp. 
40-41, 1945) agreed with this in theory but found it impractical to 
recognize a multiplicity of closely related monotypic genera of Recent 
and Pleistocene tapirs. Accordingly, he grouped all species in the 
genus Tapirus. The simplified nomenclature can be justified in this 
special case because whatever hierarchic terminology is employed in 
classification interrelationships remain the same. However, the real 
separation between each of the species should be emphasized by adding 
to Simpson’s system the available subgeneric names. 

Living and fossil tapirs were first reviewed in a classical study by 
Hatcher (Amer. Journ. Sci., ser. 4, vol. 1, art. 17, 1896). Simpson 
(op. cit.) summarized much of the information since accumulated and 
described and analyzed the osteology of North American Recent and 
Pleistocene tapirs. Concerning modern American species, these au- 
thors agreed that terrestris, bairdii, and pinchaque (roulini) are repre- 
sentative. Other named forms were regarded as either absolute syno- 
nyms or, at best, subspecies of one or another of the three species cited. 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 467 


Material 


A total of 122 specimens of Recent American tapirs were studied. 
Included in the 84 specimens examined in the U.S. National Museum 
were Tapirus terrestris (2 skins with complete skeletons, 4 skins with 
skulls, 5 complete skeletons only, 17 skulls only), Tapirus bairdia (4 
skins with skulis, 7 complete skeletons only, 41 skulls only), and 
Tapirus pinchaque (1 skin only, 3 complete skeletons only, 1 skull 
only). Included in the 38 specimens examined in the Chicago Natural 
History Museum were Tapirus terrestris (2 skins with complete skele- 
tons, 1 skin with skull, 4 skulls, skins mounted in habitat group; 21 
skulls only), Tapirus bairdii (2 complete skeletons, 6 skulls only), and 
Tapirus pinchaque (1 skin with skeleton, 1 skull only). More than 20 
skulls of the Indian tapir, Tapirus (Acrocodia) indicus, were compared 
with the American species. 

Capitalized color terms in the text are from Ridgway (Color Stand- 
ards and Color Nomenclature, 1912). 

The following abbreviations of museums are used in the lists of 
types: 

BM British Museum (Natural History) 

CNHM Chicago Natural History Museum 

MACN Museo Argentino de Ciencias Naturales ‘Bernardino Rivadavia” 
MHNP Muséum National d’Histoire Naturelle, Paris 

USNM U.S. National Museum 


Genus Tapirus Briinnich 


Distribution.—American forms are widely and nearly continuously 
distributed throughout most of the Neotropical region from about 
30 degrees south latitude in eastern Brazil to about 20 degrees north 
latitude in southern México (see map, fig. 61); altitudinal range is 
from sea level to approximately 4,500 meters above. One species, 
Tapirus (Acrocodia) indicus, is represented in Burma, Siam, French 
Indochina, the Malay States, and Sumatra. Recent tapirs are sur- 
vivors of a large family that originated in early Tertiary in the Paleo- 
arctic region and which, at successive periods of time, occupied the 
land masses now intervening between Asiatic and American species. 

Habitat and habits—Within the geographic limits defined, tapirs 
may occur in practically any wooded or grassy habitat with good 
surface supplies of water. Forests and thickets are usual daytime 
retreats, while bordering exposed areas such as grass or scrublands, 
marshes, lakes and streams with herbaceous banks, and grassy islands 
are favored nocturnal feeding resorts. Streams, whether narrow, tor- 
rential watercourses of mountain gorges or wide, sluggish rivers of 
the interior and coastal plains are indispensable refuges of all tapirs 
attacked by enemies, be they of the itch-producing, external para- 
sitic kind, or tigers, jaguars, and man. 


468 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Tapirs are omnivorous. They browse and graze. They feed on 
underwater organisms as readily as they pluck fruit, leaves, and twigs 
from shrubs and trees. They are facile of movement whether in open 
country or thick brushland, whether in water or under water. They 
are expert hill-climbers, runners, sliders, waders, and swimmers. 
Tapirs tame quickly and adjust themselves easily to artificial living 
conditions in captivity. They survive seasonal changes without ap- 
parent inconvenience when exhibited in American and European 
menageries. Ostensibly, there are no barriers in environment or 
limiting factors in tapir structure and physiology to preclude the 
coexistence of all three American species in a natural habitat suitable 
to any one of them. The Brazilian species does share a part of its 
range in northwestern South America with Baird’s tapir. Here, both 
species frequent the same feeding and watering places. On the other 
hand, the high Andean woolly tapir (Tapirus pinchaque) is not recorded 
from lower slopes of the Andes and the other tapirs are not known to 
encroach upon the upper levels, or temperate zone, of the Cordilleras. 

Characters.—External characters recorded in literature for distin- 
guishing any one of the three living American species from the others 
are, for the most part, either common to all species or are juvenal 
and individually variable characters. In all American tapirs, entire 
margin or only upper borders and lower edges of ears either conspic- 
uously trimmed or spotted with white or buff, or uniformly brown or 
black; lips edged white, gray or buff; cheeks paler than crown, the 
contrastingly paler color usually continuing onto throat, chest, and, 
to a varying degree, on belly; chin darker than cheeks and lips; upper 
parts of head and body, sides, and limbs light drab to brown in palest 
individuals, dark brown to black in darkest individuals. Juvenal 
pelage marked by a variable pattern of yellow and white spots and 
stripes covering entire body. Spotting persists past the first year of 
age and vestiges may remain (usually on limbs) in young adults. 
Apart from the urinogenital system, sexual dimorphism is not evident. 

Size.—Simpson (Bull. Amer. Mus. Nat. Hist., vol. 86, p. 77, 1945) 
calculated the following proportional differences between Pleistocene 
Tapirus excelsus and Recent T. terrestris and T. bairdii: In linear 
measurements, 7. excelsus about 1.37 times T. terrestris and about 1.19 
times JT. bairdii; in bulk and weight, 7. ercelsus over 2.5 times T. 
terrestris and about 1.7 times J. bairdzi. On the basis of these figures, 
T. bairdiit is about 1% the linear size and nearly 1% times bulkier and 
heavier than T. terrestris. The estimated difference in bulk and weight 
between the two species may be extreme. Length of skull and pro- 
portions of postcranial bones in present material confirm the greater 
average size of T. bairdii. T. terrestris averages slightly larger than 
T. pinchaque. Greatest length of skull, from gnathion to nuchal crest, 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 469 


measured on a horizontal plane, may be taken as a fair index of total 
length and bulk of any one species. This measurement, in centimeters, 
is tabulated below according to the number of functional upper molars. 
In the tabulation, the first measurement of each tooth group is average 
length of skull, figures in parentheses are extremes, and the last figure 
shows number of specimens measured. 


M! functional M2 functional M® functional 


bairdii (36 specimens) -_---_------ 39 (88-41) 10 41 (40—48) 9 42 (40—44) 17 
terrestris (41 specimens) -___---- 37 (85-41) 9 38 (86-41) 15 38 (86-41) 17 
pinchaque (5 specimens) --_--_-- 35 37, 40 37, 38 


It is practically impossible to arrive at better than a rough estimate 
of differences in size between the three living species of American 
tapirs. There may be as marked size differences between two indi- 
viduals or populations of a given species as between each of two 
“comparable” individuals or populations of different species. The 
above tabulation shows that in 7. terrestris there is no significant 
relationship between age and size after the second upper molar becomes 
functional. The same appears to be true for pinchaque, but bairdir 
continues to grow for a time after eruption of its last upper molar. 
A specimen of terrestris from Rio Grande do Sul, Brazil, with the 
largest skull, length 415 mm.,’ still has the second upper molar deep 
in the alveolus. Another skull of terrestris from Pozuzo, Peré, with 
complete and worn dentition is only 360 mm. long. Other specimens 
from the same regions indicate that difference in length between the 
two skulls is purely individual. Importance of individual variation 
in estimating size differences between species may also be demon- 
strated. A fully adult specimen of the larger bairdii, from Tiger Hill, 
C. Z., has a combined head and body length of 193 cm., while a 
“comparable” individual of the smaller terrestris from Mato Grosso, 
Brazil, measures 201 cm. 

The few available measurements indicate that the Malay tapir, 
Tapirus indicus, averages slightly larger than T. bairdivz. 


Subgenus Pinchacus* Gray 


Cinchacus [sic] Gray, Hand-list of the edentate, thick-skinned, and ruminant 
mammals in the British Museum, p. 34, 1873 (typographical error for 
Pinchacus; genotype by monotypy, Tapirus leucogenys Gray=Tapirus 
pinchaque Roulin). 


Included species—Tapirus pinchaque Roulin. 


2 Actually, in terrestris the greatest skull length, 420 mm., is of a menagerie specimen. This individual 
died at the age of 32 years after living 27 years in the National Zoological Park, Washington. 

3 The name “Cinchacus’’ originally proposed by Gray is so obviously a typographical error that it is 
quite permissible, according to article 19 of the International Code of Zoological Nomenclature, to emend 
the spelling to Pinchacus in conformity with both the etymology and the author’s intent. 


470 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Disiribution.—From upper subtropical and temperate zone forests 
into bordering ‘‘paéramos,”’ or grasslands, of the Andes in Colombia, 
Ecuador, and possibly in northern Peri and western Venezuela 
(Sierra de Mérida). (See map, fig. 61.) 

Characters—Head flattened dorsally, without mane; proboscis 
short and comparatively delicate; pelage long, coarse, thick, the 
skin well covered except usually on rump which may be marked by 
one or two patches of abraded pelage, or callouses in old adults; 
hoofs elongated, each longer than wide. Dorsal contour of skull 
flattened (fig. 58, c); median frontal line on about same horizontal 
plane as nasals and roughly parallel to horizontal plane of maxillary 
tooth row; superior parietal ridges approximating to form a low 
sagittal crest marked by a longitudinal groove (fig. 59, c) that tends 
to obsolescence in old individuals; ossification of mesethmoid carti- 
lage not extending beyond tips of nasals; exposed dorsal surface of 
maxilla as in Tapirus but may be flattened mediolaterally, maxillae 
somewhat less divergent from each other than in Tapirus; postero- 
lateral maxillary process as in Tapirus, overlying frontal process and 
not entering into composition of inner lateral wall of narial meatus; 
posterolateral margin of premaxilla slightly, entirely, or not at all 
embraced laterally by outer anterior border of maxilla; nasals as in 
Tapirus but more elongate, descending process more delicate. First 
upper premolar with or without cinguloid shelf extending anteriorly 
from protocone (fig. 60, a, 6); upper incisors distinctly opisthodont. 

Remarks.—Cranial contours of the mountain tapir show least 
departure from primitive lines; cranial characters generally, and 
particularly those associated with the comparatively little-developed 
proboscis, are less specialized than in other Recent species; dentition, 
as manifested by the first upper premolar, is variable. Ecuadorian 
specimens show the simple condition, with cinguloid shelf absent, 
while the only authentic Colombian skull examined shows the pre- 
molar as in true Tapirus. 

Distribution of Pinchacus points to its prior arrival into South 
America and at a period when a temperate climate prevailed at sea 
level in equatorial latitudes. It inhabits an area representing part 
of the original Colombian Central Land Mass, the South American 
side of the intercontinental land bridge where Tertiary mammals 
entering from North America established foothold. Present restric- 
tion of Pinchacus to the Colombian Central Land Mass, now the 
temperate zone of the bulk of the Venezuelan, Colombian, and Ecua- 
dorian Andes, probably is the result of an inherited urge for susten- 
tation in cooler climates pari passu with increasing rise in height 
of the Andes above sea level and rising temperatures at sea level. 
Newly established tropical zone habitats at the base of the Andes 
were invaded subsequently by other kinds of tapirs. 


MAMMALS OF NORTHERN COLOMBIA—-HERSHKOVITZ 471 
Tapirus pinchaque Roulin 


(Woolly, Andean, or mountain tapir, ‘“danta lanuda’” or ‘danta 
cordillerana’’) 


Le Pinchaque Cuvier, Ann. Sci. Nat., Paris, ser. 2 (Zool.), vol. 17, p. 110 (pp. 
107-112), 1829 (advance notice with description and comparisons of the 
“pinchaque” of Roulin). 

Tapir pinchaque Roulin, Ann. Sci. Nat., Paris, ser. 2 (Zool.), vol. 18, p. 46, pls. 
1, 2, figs. 1-3, 1829.—Goudot, Compt. Rend. Acad. Sci., Paris, vol. 16, 
p. 331, 1843 (Las Juntas, Rfo Combeima, southern foot of Mt. Tolima, 
Colombia, altitude, 1,918 meters; description, habits; tracks seen between 
1,400 and 4,400 meters above sea level).—Blainville, Ostéographie, text, 
vol. 4, fasc. 19, p. 22, 1846 (osteology). 

Tapirus pinchaque, Hunter, The natural history of the quadrupeds of Paraguay 
. .. (translated from the Spanish of Felix de Azara), vol. 1, p. 113, 1838, 
Edinburgh (ref.). 

Tapirus pinchacus [sic], Gray, Proc. Zool. Soc. London (1867), p. 884, 1868; 
ibid., (1872), p. 484, 1872 (original description quoted; bibliography; syn- 
onymy).—Blainville, Ostéographie, text, vol. 4, fasc. 19, p. 51, 1846 (“‘7. 
roulini Fischer,’”’ in synonymy), atlas, vol. 4, pl. 3, 1846 (skull of type; 
skull of specimen collected by Goudot in Colombia).—Déderlein, Ueber das 
Skelett des Tapirus Pinchacus, Inaugural-Dissertation, 1877 (osteology; 
distribution). 

Tlapirus] Roulinii Fischer, Synopsis Mammalium, Addenda, p. 604 [sic=406], 
1830 (new name for Tapir pinchaque Roulin). 

Tapirus roulini [sic], Sclater, Proc. Zool. Soc. London (1870), p. 51 and footnote, 
1870 (report of specimen seen by Robert A. White on Volcdn de Puracé, 
Cordillera Central, Colombia, altitudinal range reported as between 3,500 
and 4,200 meters above sea level; ‘‘the first Latin name applied to this tapir 
appears to be roulini [sic] Fischer’’); ibid., (1872), p. 604, fig. 2 (masals), 
1872 (skull collected by Buckley, undoubtedly in Ecuador).—Hatcher, 
Amer. Journ. Sci., ser. 4, vol. 1, p. 173 and text, pl. 4, figs. 2, 2a, pl. 5, fig. 2, 
1896 (cranial characters, comparisons, phylogeny, synonymy: Pinchacus, 
villosus, leucogenys, aenigmaticus [=terrestris], pinchaque).—Lydekker, Cata- 
logue of the ungulate mammals in the British Museum (Natural History), 
vol. 5, p. 44, 1916 (synonymy: pinchaque, villosus, leucogenys, enigmaticus 
[sic=terrestris]; specimens from Sufiac and ‘‘Assuay,’’ Ecusdor).—Simpson, 
Bull. Amer. Mus. Nat. Hist., vol. 86, pp. 39-80, pl. 10, fig. 2, 1945 (cranial 
and dental characters, comparisons, phylogeny).— Crandall, Animal kingdom, 
Bull. New York Zool. Soc., vol. 44, No. 1, 1951 (description of live animal in 
captivity). 

Tapirus roulinii, G. M. Allen, Extinct and vanishing mammals of the Western 
Hemisphere, p. 404, 1942 (part, not description; Department of Santander, 
Colombia, 8,000—10,000 feet altitude; Cordillera de Llanganates, Cordillera 
Oriental, Ecuador, 14,000 feet altitude). 

Rhino {choerus] villosus Wagler, Natiirliches System der Amphibien . . ., footnote 
2, p. 17, 1830 (name proposed for “le Pinchaque’’ Cuvier= Tapir pinchaque 
Roulin). 

Tapirus villosus Wagner, in Schreber, Die Saéugthiere in Abbildungen nach der 
Natur, mit Beschreibungen, vol. 6, p. 392, pl. 319b, 1835 (new name for 
Tapir pinchaque Roulin).—Tschudi, Untersuchungen iiber die Fauna Peruana, 
Therologie, pp. 213, 215, 1844 (description, distribution in Pert). 


472 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Tlapirus] andicola Gloger, Gemeinniitziges Hand- und Hilfsbuch der Natiir- 
geschichte .. ., vol. 1, p. 124, 1842 (new name for pinchaque Roulin). 

Tapirus leucogenys Gray, Proc. Zool. Soc. London (1872), p. 488, pl. 21, 1872 
(type locality, ‘‘“Ecuador, on the Cordilleras at Sunia and Assuay’’).— 
Lydekker, Catalogue of the ungulate mammals in the British Museum 
(Natural History), vol. 5, p. 44, 1916 (type from “‘Assuay”’; listed as synonym 
of rouliniz). 


Types—Of pinchaque Roulin, adult male, skull only, MHNP, 
collected by Roulin; of lewcogenys Gray, adult male lectotype (desig- 
nated by Lydekker, supra cit.), skin and skull mounted, BM 72.1.24.3-4 
(1577 b), collected by Clarence Buckley. 

Type localities —Of pinchaque Roulin, Péramo de Sumapaz, Cordil- 
lera Oriental, south of Bogoté, extreme southern part of Department 
of Cundinamarca, Colombia; of leucogenys Gray, Péramo del Azuay, 
Cordillera Oriental, southern Ecuador (restricted by lLydekker, 
supra cit.). 

Distribution —Generally as for the subgenus; actual specimens 
recorded in scientific literature were taken only in the Cordilleras 
Oriental and Central of Colombia and the Cordillera Oriental of 
Ecuador, from 5 degrees north latitude (Mt. Tolima) to about 4 
degrees south latitude (Azuay) (see map, fig. 62); altitudinal range 
between 2,000 and 4,400 meters. Carriker (in G. M. Allen, supra cit.) 
reported tapirs common at altitudes from 8,000 to 10,000 feet in the 
Cordillera Oriental, Department of Santander, Colombia, at the 
Venezuelan border, about 7 degrees north latitude. No signs of 
T. pinchaque were seen by the writer in the comparatively low Sierra 
de Perij4, the extension of the Cordillera Oriental north of Santander, 
and the species is unknown in the Colombian Cordillera Occidental. 
Tschudi (supra cit., p. 215) recorded the woolly tapir from the upper 
forest zone of the Peruvian Andes at elevations between 7,000 and 
8,000 feet. This report, never since confirmed, was based on testi- 
mony of natives who killed the animal in “mittleren Peru in der Ceja 
von Comas, Huancavelica etc.’”? Published records of tapir tracks 
noted in the Andes at elevations in the neighborhood of 1,500 meters 
above sea level may refer to any species, although no museum speci- 
mens of tapir have actually been collected anywhere within 300 meters 
of this altitude. 

Characters—Those given for the subgenus. Some general external 
characters are mentioned under the generic heading. Skin of adult 
female from Rio Majuas, Colombia, collected by the writer in 1951, 
is blackish brown on back, sides of body, upper parts of limbs, and 
tail; pelage of rump on either side of middorsal line abraded; individual 
cover-hairs of dorsum 1 to 1 centimeters long, with very fine, crinkly, 
brown tips, black basally; hairs on sides of body and chest become 
progressively longer to approximately 3 cms. with basal portions 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 473 


brown; long, crinkly wool-hairs sparsely present on sides and under- 
parts; guard-hairs scattered, slightly stiffer than cover-hairs and 1 to 
2 cms. longer; hairs of lower parts of limbs short, harsh, black with 
fine brown tips. There is a thin sprinkling of white hairs over all dark 
parts of body. Head is blackish brown dorsally, with whorl on fore- 
head ; ear well haired on outer side, anterior and upper borders blackish 
brown, behind brown sprinkled with whitish; inner side of ear prac- 
tically bare except for a thin concentration, not at all conspicuous, of 
whitish hairs along borders. Sides of head and neck are brownish, 
the hairs with gray bases; muzzle blackish brown with tip and sides 
strongly grizzled and sharply demarcated from narrow white band 
encircling mouth; underparts blackish brown, pelage of throat long, 
somewhat matted, that of belly shorter, and directed forward; anal 
region thinly haired, whitish. A pair of teats is present. 

Another adult female exhibited in the New York Zoological Park is 
described by Crandall (Animal Kingdom, Bull. New York Zool. Soc., 
vol. 54, p. 3, figs., 1951) as “clothed [on body] with dense, matted 
hair, blackish brown in color. . . head paler. Individual hairs 
from the back are approximately an inch long, most of them with one 
or two kinks. The white fringe at the ear tips, present in all tapirs, 
is especially conspicuous in Panchita, because of the length and 
density of the hair. . .. Also, the eyes are pale brown—not blue 
as often stated.” 

Skin of a third adult‘ is blackish brown on back, hairs black 
terminally, dark brown basally, the brown portion increasing pro- 
gressively toward posterior end of back; sides mixed blackish and 
auburn, becoming nearly uniformly Tawny on belly, chest, and 
posterior sides of thighs; limbs blackish brown with a scattering of 
buffy and ochraceous hairs; rump with a nearly bald area; top of head 
dark reddish brown, nearly black; muzzle brown, approximately 
auburn; sides of face mixed brown, ochraceous and buffy; upper lip 
and chin buffy, throat brown; ears auburn edged with buffy to 
ochraceous. 


4 Sex unknown, it was removed from exhibition in the halls of the U. S. National Museum in March, 
1911, and preserved as skull and two flat pieces of hide. Somehow, the skin was given the same catalog 
number (USNM 61221) as one of two skins of 7. bairdii formerly mounted and exhibited by the 
Gustemalan Commission in the Columbian Exhibition. The skull was likewise misnumbered but in 
September 1938 was renumbered 267894. Most likely, the specimen originated in Ecuador but it may 
have come from Colombia. According to records of the Smithsonian Institution, Gen. 8. A. Hurlbut 
donated skins with complete skeletons of two mountain tapirs he secured in Tolima, Colombia. The 
skeletons were entered in the Mammal Division “bone” catalog on November 11, 1871, as numbers 11883 
and 11884. There is no account of corresponding skins, if ever accessioned. In 1872, President Garcia 
Moreno of Ecuador presented through the U. S. Minister to Ecuador, the Hon. E. Rumsey Wing, a 
complete skeleton of mountain tapir. The specimen is entered in the ‘‘bone’”’ catalog as No. 12759. How- 
ever, according to the Annual Report of the Smithsonian Institution for 1873 (p. 47), a mounted specimen 
was also donated by President Garcfa Moreno. A “skin” catalog entry in 1874, No. 11869, is of a 
mounted “Tapirus Roulini’’ without further data, but almost certainly the specimen now at hand. 


279127—54——2 


474. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


. Measurements.—Of the type specimen, fully adult male with worn 
dentition (ex Roulin, supra cit., p. 32): Total length, 5 ft., 6.5 in., 
in old French system, or approximately 1,800 mm.; height at withers, 
2 ft., 9 in., or about 900 mm.; length of forefoot, 1 ft., 4 in., or about 
433 mm.; hind foot slightly shorter; circumference of upper foreleg, 
over 16 in.; circumference of upper hind leg considerably less; weight, 
completely eviscerated, between 240-250 French pounds. Of the 
fully adult female, from Rio Majuas, upper Rio Magdalena Valley, 
Huila, Colombia, 2,700 meters altitude: Head and body, about 
2,000 mm.; tail, 50 mm.; hind foot (approx., from skeleton) 310 mm.; 
ear, 170 mm.; height at withers (approx., from skeleton) 870 mm.; 
height at rump (approx., from skeleton) 910 mm.; weight, 583 pounds 
(including fetus of approximately 10 pounds). Of ‘‘Panchita,’’ 
adult female, on 21 November, 1951, age between 3% and 4 years, 
measurements according to Lee S. Crandall, General Curator of the 
New York Zoological Park (in litt.): “Tip of nose to base of tail, 
6 ft. [1,829 mm.]; tail, 2 in. [51 mm.]; height at shoulder, 2 ft., 7% in. 
[80 mm.]; height at hips 2 ft., 944 in, [851 mm.]; greatest circumference, 
4 ft., 3 in. [1,295 mm.].” At reported age of 2% years, Panchita weighed 
223 pounds. 

Remarks.—The abraded, bald, or calloused areas of the rump are 
evidently consequences of the tapir’s habit of sitting or reclining on 
either side of its rump and from scrubbing the affected areas against 
rocks, gravelly ground, and tree trunks to relieve itching. The large 
size and quantity of ti¢ks attached to all species of tapirs is notorious. 
Similar abrasions of the rump in short and thinly haired species 
living in tropical lowlands where rocks are rare could easily escape 
detection. Goudot (Compt. Rend. Acad. Sci., Paris, vol. 16, p. 331, 
1843) reports that native hunters attribute the peculiarities of the 
rump to the animal’s proclivities for sliding downhill in, presumably, 
a sitting posture. This explanation need not be given serious con- 
sideration. No bald spot or abrasion appeared on the rump of the 
young adult female tapir killed by Goudot in the Colombian Cordil- 
lera Central and none was present on the type of 7. lewcogenys Gray. 
On the other hand, abrasions mark the rumps of the specimens 
described in the preceding section, and were present in two live 
woolly tapirs that arrived at the New York Zoological Park (see 
below). Both individuals were hand raised from the very young, 
spotted-and-striped stage. Neither of them had been exposed to 
conditions of life conducive to downhill rump sliding. 

Color of upper border of ear is variable. The female from Majuas, 
Colombia, described above and the two males observed by Roulin 
lack white on upper borders of the ears. Ears of the type of leucogenys 
Gray are described as “with scarcely any indication of white edges.” 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 475 


All other specimens of woolly tapirs of which external characters 
are known and recorded or described herein have white ear tips. 

In skinning the woolly tapir taken in Colombia, the writer was im- 
pressed by its comparatively thin hide. Hides of the Brazilian, 
Baird’s, and Indian tapirs are notoriously thick and range in thickness 
in the order named, that of the last the heaviest. Characteristic extra 
thickness of hide of nape in these species is altogether absent in the 
maneless T. pinchaque. The Indian tapir with maximum thickness of 
hide offers a deep armor where most vulnerable to the tiger’s fangs. 
Hide of nape combined with mane in Brazilian and Baird’s tapirs are 
equally good fenders against the smaller-toothed and weaker-jawed 
jaguar. Mountain fastnesses of the woolly tapir are rarely, if ever, 
visited by jaguars. Mountain lions (Felis concolor) occur within the 
range of T. pinchaque but rarely attack any wild animal larger than 
a brocket (Mazama). Bears (Tremarctos ornatus) sometimes prey 
on woolly tapirs but their technique of attack, distinct from that of 
cats, gives no special advantage to quarry with a well-protected nape. 

Tapirs eat anything edible and a surprisingly large amount of ma- 
terial such as woody twigs and mud, which is not generally regarded 
as edible. However, in any given habitat the bulk of the tapir’s diet 
consists of green shoots of the commonest browsing plant. Stomachs 
of woolly tapirs examined by the writer, by Roulin, and by Goudot 
(supra cit.) contained mostly ferns and shoots of “‘chusque,”’ a, trailing 
bamboo of the genus Chusquea. These are the dominant plants in 
many situations of the steeper, more sterile wooded slopes of the 
Andean temperate zone. In “péramo” zone, according to Goudot 
(p. 334), woolly tapirs eat tender shoots of ‘“‘frailej6n’”’ (Espeletia) as 
well as those of rough grasses. 

The word “pinchaque” is, according to Roulin, the name of a large 
fabulous animal believed to live within the Colombian range of the 
woolly tapir. It may refer to the extinct Mastodon but few, if any, 
Colombians now use the term. ‘‘Danta” is the name applied indis- 
criminately by natives to all three species of tapirs. The terms woolly 
tapir, mountain tapir, and Andean tapir have been in use in the Eng- 
lish language for over a century. Their Colombian equivalents are 
“danta lanuda” and “danta cordillerana.”’ Ecuadorians of the east- 
ern slope of the Cordillera Oriental distinguish the woolly from the 
Brazilian tapir by the names “‘danta negra’”’ and “danta café,’’ respec- 
tively. ‘“Huagra” is the Quechua term for tapir. For lack of another 
word in their vocabulary, Quechua-speaking Indians of Ecuador use 
“huagra” also for domestic cattle. 

The first specimen of Tapirus pinchaque to be exhibited alive out- 
side its country of origin is ‘‘Panchita,’”’ received in the fall of 1950 
by the New York Zoological Society’s park in the Bronx, New York. 
The animal, a female, was approximately 2% years of age at the time. 


476 PROCEEDINGS OF THE NATIONAL MUSEUM Vor. 103 


According to Crandall (Animal Kingdom, Bull. New York Zool. Soc., 
vol. 44, no. 1, pp. 3-8, 1951), the specimen, from the Cordillera 
Oriental of Ecuador, lived “‘as a village pet in a hamlet called Borja, 
sixty miles |kilometers is surely intended] to the east of Quito at an 
elevation of about 6,000 feet... . ‘Panchita’, as the villagers 
called the animal, had been captured higher up the Andean range 
while still in her striped-and-spotted coat of tapir infancy.’’ The 
second captive tapir, a male from Papallacta, east of Quito, Ecuador, 
about 3,150 meters above sea level on the eastern slope of the Cordil- 
lera Oriental, was received June 19, 1952, by the same zoological park 
(cf. Animal Kingdom, Bull. New York Zool. Soc., vol. 45, Nos. 1, 2, 
1952). It had been sent by Charles Cordier, the dealer who also 
secured the first live tapir. 

Contrary to all ruling, previous authors have attempted to replace 
the first valid name, pinchaque Roulin, with Latin names of which the 
earliest and most commonly cited is rowlinit Fischer. The specific 
name pinchaque was properly proposed in combination with the 
generic synonym Tapir as technical name for the woolly tapir. 
Roulin explicitly borrowed pinchaque from the vernacular term 
applied by some Colombians to a large legendary animal, possibly 
the extinct mastodon. In an advance notice of the discovery of the 
woolly tapir, Cuvier (supra cit.) discreetly used ‘‘pinchaque”’ strictly 
as the vernacular term for the new species, leaving the formal proposal 
of a technical name to Roulin. 

Published descriptions and figures of misidentified species of 
American tapirs have led to some garbled accounts in current litera- 
ture. External characters attributed to the woolly tapir by G. M. 
Allen (supra cit.) are derived from a figure and description by Sclater 
(Proc. Zool. Soc. London (1878), p. 631, pl. 39, 1878) of a living ex- 
ample of T. terrestris exhibited in the London Zoological Gardens. 
The individual was first misrepresented as ‘‘Tapirus roulint.” Later, 
as the result of a post mortem, Sclater (ibid. (1885), p. 718, 1886) 
discovered his error and emended the name to Tapirus americanus 
(=T. terrestris). Tapirs secured by Buckley at Sarayacu, eastern 
Ecuador, and mentioned by G. M. Allen are also misidentified repre- 
sentatives of TJ. terrestris (cf. antea, in synonymy of T. terrestris 
terrestris). 

An early revision of tapirs by Gray (Proc. Zool. Soc. London (1872), 
pp. 483-492, pls. 21, 22, 1 fig., 1872) has been a popular but confusing 
and misleading source of information. The work is characterized by 
numerous typographical errors, misquotations of authors, contra- 
dictions, and assumptions derived from specimens mislabeled as to 
sex and locality and mismatched as regards skins and corresponding 
osteological material. Gray’s description of external characters of 
the adult leuwcogenys may be that of terrestris, but the skull, as analyzed, 


MAMMALS OF NORTHERN COLOMBIA—-HERSHKOVITZ 477 


certainly pertains to pinchaqgue. It may be necessary to restrict the 
type of leucogenys to the skull only. The young individual described 
by Gray as ‘“‘Tapirus leucogenys jun., or T. aenigmaticus,” if correctly 
figured (pl. 21, and skull, p. 491) must be regarded as T. terrestris. 
The skin of this specimen may be that of a mountain tapir but the 
skull as figured by Gray is unquestionably that of a lowland tapir. 
Indeed, Gray (pp. 484, 491, 492) questioned both locality data and the 
association of skin and skull as given by the collector, Clarence 
Buckley. Present determination of types and type localities of 
Gray’s specimens follow Lydekker (Catalogue of the ungulate mam- 
mals in the British Museum (Natural History), vol. 5, p. 44, 1916). 

In addition to specimens examined by the writer, there is a mounted 
skin and skeleton and a skull only from Ecuador in the Museum 
of Comparative Zoology and two specimens from the Cordillera de 
los Llanganates (Cordillera Oriental), Ecuador, in the Academy of 
Natural Sciences of Philadelphia. The species is also poorly repre- 
sented in Kuropean museums. 

Specimens examined.—Six. Corompra: Huila, San Agustin, Rio 
Majuas, upper Rio Magdalena, 2,700 meters, 1 female, skin and 
skeleton (CNHM); no precise locality, 2 complete skeletons (USNM). 
Ecvuapor: Cordillera de los Llanganates, upper Rio Pastaza drainage, 
Cordillera Oriental, 1 skull (CNHM); no precise locality, 1 complete 
skeleton (USNM). No locality (probably Ecuador, possibly Colom- 
bia), 1 skin with skull (USNM). 


Subgenus Tapirus Briimnich 


Tapirus Brinnich, Zoologiae fundamenta ..., pp. 44, 45, 1772.—Scopoli, J. A- 
Scopoli . . . Introductio ad historiam naturalem ..., p. 492, 1777.— 
Merriam, Science, new ser., vol. 1, p. 376, 1895 (based on ‘‘Le Tapir’’ or 
Tapirus Brisson, Regnum Animale, 1762). 

Tapir Blumenbach, Handbuch der Natiirgeschichte, ed. 1, vol. 1, p. 129, 1779 
(genotype, by monotypy, Tapir suillus Blumenbach=Tapirus terrestris 
Linnaeus).—Zimmermann, Geographische Geschichte ... , vol. 2, p. 154, 
1780 (genotype, by monotypy, anta Zimmermann=+¢errestris Linnaeus) .— 
Gmelin, Caroli a Linné systema naturae .. . , ed. 13, vol. 1, p. 216, 1788 
(genotype, by monotypy, americanus Gmelin=tapir Erxleben=terrestris 
Linnaeus). 

Syspotamus Billberg, Synopsis faunae Scandinaviae, vol. 1, pt. 1, Mammalia, 
Conspectus 4 (before p. 1), 1827 (new name for Tapir Gmelin). 

Rhinochoerus Wagler, Natiirliches System der Amphibien ..., p. 17, 1830 
(proposed for Tapirus Brisson). 

Tapyra Liais, Climats, géologie, faune et géographie botanique du Brésil, p. 397, 
1872 (emendation of Tapirus). 


Included species—Tapirus terrestris Linnaeus. 


Distribution.—Tropical zones of mainland South America, from 
Rio Grande do Sul, Brazil, and the Chaco of Argentina, Paraguay, 


478 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


and Bolivia, north through the Amazonian regions of Brazil, Bolivia, 
Pert, Ecuador, and Colombia; in the Guianas and Venezuela west 
across the Sierra de Perija of the Cordillera Oriental into northern 
Colombia at least as far as the Rfo Atrato (see map, fig. 61); unknown 
from the Pacific coastal plains of South America; altitudinal range 
from sea level to 1,200 meters above, not certainly recorded from 
higher altitudes. 

Occurrence of the Central American Tapirella in northwestern 
South America presupposes the probable existence of typical Tapirus 
in Central America. However, bonafide records of T. terrestris from 
Central America do not exist. A specimen of 7. terrestris, supposedly 
from Talamanca, Costa Rica, was recorded by Goldman (Smithsonian 
Misc. Coll., vol. 69, p. 83, 1920). Later authors have cited the same 
record. The specimen in question, a skull only, was entered into the 
catalog of the mammal collection of the U. S. National Museum in 
1873 or 1874 along with other skulls of terrestris and pinchaque and 
14 skulls of bairdit from Costa Rica. No doubt, the skull, without 
original locality data, was inadvertently included in the catalog with 
the Costa Rican material of Prof. Gabb. 

Characters.—Crest of head rising abruptly from behind base of muz- 
zle and surmounted by a low, narrow mane that continues back to 
withers, sometimes to behind middle of back; proboscis well developed, 
fairly bulky; pelage short, stiff, and usually not completely hiding 
skin; hoofs short, broad, the middle always wider than long. Dorsal 
contour of braincase (fig. 58, a) decidedly convex, the median frontal line 
rising steeply from horizontal plane of nasals and diverging even more 
sharply from horizontal plane of maxillary tooth row; superior ridges 
of parietals united to form a single arched sagittal crest (fig. 59, a); 
ossification of vertical mesethmoid cartilage usually not extending 
anteriorly beyond middle of nasals, rarely to tips of nasals; maxillae 
divergent, exposed dorsal surface of each rounded mediolaterally, 
without rising process in front of plane of infraorbital foramen; 
posterolateral maxillary process overlying anterior frontal process and 
not entering into composition of inner lateral wall of narial meatus; 
posterolateral margin of premaxilla forming a nearly straight or a 
slightly curved line, not embraced laterally by anterior margin of 
maxilla; nasal with thick descending sigmoid process overlapping 
maxilla. First upper premolar with cinguloid shelf extending an- 
teriorly from internal cusp (fig. 60, @); upper incisors slightly 
pro-odont. 

Remarks.—A glance at the striking sagittal crest of the skull or 
the head of the living animal is sufficient for distinguishing 7’. ter- 
restris from all other Recent tapirs. The crest appears to be an 
extreme development of a condition that is incipient in 7. pinchaque 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 479 


where the parietal ridges almost or barely unite to form a low, 
inconspicuous crest. Cranial characters associated with the proboscis 
are, in 7’. terrestris, hardly distinguishable from those of 7. pinchaque. 
Nevertheless, the proboscis of the Brazilian tapir is more developed. 


Tapirus terrestris terrestris Linnaeus 


(“Tapir” of Brazilian Tupi; ‘“danta,” ‘“anta,” and “gran bestia’ of 
Spanish and Portuguese; “‘huagra” of Quechua Indians of Peruvian 
and Ecuadorian Amazonas, ‘“mborebi” of Paraguayan Guaranf; 
‘“maipuri” or “manipuri’”’ of Guianan Gahibi and Macusi Indians; 
“bushcow”’ of British Guianan colonists) 


Hippopotamus terrestris Linnaeus, Systema naturae, ed. 10, vol. 1, p. 74, 1758. 

Hydrochaerus tapir Erxleben, . . . Systema regni animalis. .. . Mammalia, p. 
191, 1777 (range given: Istmo de Darién to Rio Amazonas). 

Tapir suillus Blumenbach, Handbuch der Naturgeschichte, ed. 1, vol. 1, p. 129, 
1779 (based on Allamand suppl. in Buffon, Histoire naturelle, vol. 15, Holland 
ed., pls. 9, 10, of male and female T. terrestris exhibited in Amsterdam, origin 
unknown, possibly Surinam). 

Tapir (Anta) Zimmermann, Geographische Geschichte . . . vol. 2, p. 154, 1780, 
part (based primarily on the tapir of Marggraf and Linnaeus=terrestris 
Linnaeus). 

Tapir americanus Gmelin, Caroli a Linné. . ., Systema naturae, ed. 13, vol. 1, 
p- 216, 1788 (primary reference, Brisson, Regnum animale, p. 119, 1756, the 
tapir of Guiana and Brazil; range given: Istmo de Panama to Rio Amazonas). 

Tlapirus] americanus, Tschudi, Untersuchungen iiber die Fauna Peruana, Therol- 
ogie, pp. 213-215, 1844 (Perd; description, habits and distribution). 

Tapirus rufus, Fischer, Zoognosia tabulis synopticis illustrata . . ., vol. 3, p. 292, 
1814 [ed. 3] (Guiana). 
Tapir maypuri Roulin, Ann. Sci. Nat. Zool. Paris, vol. 18, p. 45, 1829 (name 
derived from the vernacular of Indians living between right bank of the 

Caroni and mouth of Amazon). 

Tapirus laurillardi Gray, Proc. Zool. Soc. London (1867), p. 881, figs. a, B, 1-4, 
1868. 

Tapirus terrestris laurillardi, Lydekker, Catalogue of the ungulate mammals in 
the British Museum (Natural History), vol. 5, p. 43, 1916 (Venezuela?). 
Tapirus leucogenys, juv., Gray, Proc. Zool. Soc. London (1872), p. 490, pl. 22, 

fig. 1 (animal), fig. p. 491 (skull), 1872. 

Tlapirus] aenigmaticus Gray, Proc. Zool. Soc. London (1872), p. 490, pl. 22, fig. 1, 
1872 (alternative name for “‘lewcogenys, juv.’’ Gray).—Lydekker, Catalogue 
of the ungulate mammals in the British Museum (Natural History), vol. 5, 
p. 44, 1916 (type from ‘“‘Sunac,” synonym of “‘roulini’’). 

Tapirus ecuadorensis Gray, Proc. Zool. Soc. London (1872), p. 492, pl. 22, fig. 2, 
1872.—Thomas, Proc. Zool. Soc. London (1880), p. 400, 1880 (synonym of 
roulinit). 

Tapirus (terrestris) peruvianus Gray, Proc. Zool. Soc. London (1872), p. 624, pl. 
45, 1872. 

Tapyra sabatyra Liais, Climats, géologie, faune et géographie botanique du Brésil, 
p. 397, 1872 (name suggested as more ‘‘appropriate’’ for the Brazilian tapir) . 

Tapirus pinchaque ou roulinii . . . variété brasiliensis, Liais, Climats, géologie, 
faune et géographie botanique du Brésil, p. 398, 1872. 


480 PROCEEDINGS OF THE NATIONAL MUSEUM. VOL. 108 


Tapirus tapirus Merriam, Science, new ser., vol. 1, p. 376, 1895 (new combination 
of tapirus Brisson plus fapirus Merriam ex Brisson=terrestris Linnaeus). 
Tapirus americanus Brisson, var. mexianae Hagman, Archiv Rass.-Ges. Biol., vol. 

5, p. 22, fig. 1, 1908. 

Tapirus terrestris mexiana, Lydekker, Catalogue of the ungulate mammals in the 
British Museum (Natural History), vol. 5, p. 48, 1916. 

Tapirus spegazziniti Ameghino, Anales Mus. Nac. Buenos Aires, vol. 20, p. 31, 
pls. 5, 6, December 1909 (separate) ; ibid., ser. 3, vol. 18, p. 31, pls. 5, 6, 1911 
(bound volume). 

Tapirus terrestris spegazzinii, Lydekker, Catalogue of the ungulate mammals in 
the British Museum (Natural History), vol. 5, p. 43, 1916. 

Tapirus terrestris guianae Allen, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 566, 1916. 

Tapirus anulipes Hermann, Mitt. Zool. Mus. Berlin, vol. 11, pt. 1, p. 167, 1924. 

Tapirus (Hippopotamus) terrestris, Hatcher, Amer. Journ. Sci., vol. 1, p. 174, pl. 
4, figs. 1, 1a, pl. 5, fig. 1, 1896 (cranial characters; comparisons; phylogeny; 
synonymy: americanus, anta, suillus, laurillardi, equadorensis [sic]). 

Tapirus terrestris, Gray, Proc. Zool. Soc. London (1867), p. 879, 1868 (revision; 
skulls only from Brazil; Berbice and Demerara in British Guiana.)—Sclater, 
Proc. Zool. Soc. London (1873), p. 193, fig. p. 194, 1873 (Paraguay).— 
Osgood, Field Mus. Nat. Hist. Publ., Zool. Ser., vol. 10, p. 46, 1912 (Empa- 
lado Savanas, Zulia, Venezuela)—Lydekker, Catalogue of the ungulate 
mammals in the British Museum (Natural History), vol. 5, p. 41, 1916 
(synonyms: tapir, anta, americanus, sutllus, ecuadorensis, tapirus Palmer 
ex Brisson; type locality, Brazil or Paraguay).—Goldman, Smithsonian 
Mise. Coll., vol. 69, p. 83, 1920 (skull only in U. 8. National Museum labeled 
“Talamanca, Costa Rica’”’).—Tate, Bull. Amer. Mus. Nat. Hist., vol. 76, 
p. 222, 1939 (Duida and Auydn-tepuf, Venezuela; British Guiana).—Simp- 
son, Bull. Amer. Mus. Nat. Hist., vol. 86, p. 42, tables 1-8, figs. 1-4, 
pl. 10, fig. 2, 1945 (osteological and dental characters; comparisons; phy- 
logeny; Mato Grosso, Brazil; Colombia; eastern Peri; British Guiana; 
“South America”’)—Goodwin, ibid., vol. 87, p. 451, 1946 (‘‘Talamanca, 
Costa Rica’). 

Tapirus terrestris terrestris, Lydekker, Catalogue of the ungulate mammals in 
the British Museum (Natural History), vol. 5, p. 42, 1916 (Rio de Janeiro; 
Taquara, Rio Grande do Sul, Brazil; ‘Brazil’; Berbice and Demerara, 
British Guiana; Surinam). 

Tapirus roulini [sic], Sclater (nec Fischer), Proc. Zool. Soc. London (1878), p. 631, 
pl. 39, 1878.—Thomas, Proc. Zool. Soc. London (1880), p. 400, 1880 (Sara- 
yacu, Ecuador; striped juvenals without skulls; part of original series de- 
scribed by Gray as ecuadorensis placed in synonymy of rouliniz).—Lydekker, 
Catalogue of the ungulate mammals in the British Museum (Natural His- 
tory), vol. 5, p. 44, 1916 (part; enigmaticus [sic] in synonymy only). 

Tapirus roulini [sic, nec Fischer] x Tapirus americanus, Sclater, Proc. Zool. Soc. 
London (1882), p. 311, 1882 (specimen of “‘roulini’”’ same as the “roulinz” of 
Sclater, 1878, supra cit.). 

Tapirus dowii (?), Sclater (nec Gill), Proc. Zool. Soc. London (1882), p. 391, pl. 
23, 1882 (Rfo Yuruari, Venezuela). 

Tapirus americanus, Sclater, Proc. Zool. Soc. London, 1885, p. 718, 1886 (correc- 
tion of identification of “‘roulini” Sclater, 1878, supra cit., and “‘dowii (?)” 
Sclater, 1882, supra cit.)—Déderlein, Ueber das Skelett des Tapirus pin- 
chacus; Inaugural-Dissertation, p. 3, Bonn, 1877 (distribution). 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 48]. 


Types.—Of terrestris Linnaeus, tapir Erxleben, anta Zimmermann, 
americanus Gmelin, suillus Blumenbach, none in existence, names 
based on bibliographic references; of rufus Fischer, skin and skull, 
MHNP; of maypuri Roulin, sabatyra Liais, none in existence, names 
proposed as substitutes for terrestris Linnaeus; of brasiliensis Liais, 
none in existence, name based on reported differences alleged to exist 
in a “variety’’ of Brazilian tapir; of /aurillardi Gray, adult, skull only, 
BM 52.12.9.3. (709 g), purchased from Brandt of Hamburg in 1852; 
of aenigmaticus Gray, immature, skull only if skin mismatched, BM 
72.1.24.9-10 (1577 f), collected by Clarence Buckley; of ecuadorensis 
Gray, striped juvenal, male, skin mounted, BM 72.1.24.13, collected 
by Buckley, original number, 13; of peruvianus Gray, striped juvenal, 
skin and skull, presumably the specimen later listed by Gray (Hand- 
list of the Edentate, Thick-skinned and Ruminant Mammals in the 
British Museum, p. 33, 1873) as BM 69.3.31.9. (38 d), with skull, 
72.4.11.4 (709 1), collected May 31, 1868, by E. Bartlett; of mezxianae 
Hagmann, young adult, skull only, ‘Zoological Collection, Strass- 
burg”; of spegazzinit Ameghino, adult male, skull only, MACN, col- 
lected by Carlos Spegazzini; of guianae Allen, young adult, skull 
only, AMNH 36198, collected by Leo E. Miller; of anulipes Her- 
mann, young adult with persistent juvenal streaks and spots, 
especially on limbs, observed in zoological garden in Corumb4é, Mato 
Grosso, Brazil. 

Type localities —Of terrestris Linnaeus, Brazil, restricted to Pernam- 
buco by Thomas (Proc. Zool. Soc. London (1911), p. 155, 1911); of 
anta Zimmermann, here restricted to Pernambuco; of sabatyra Liais, 
same as for terrestris Linnaeus; of laurillardi Gray, ‘“‘SSouth America,” 
here restricted to Pernambuco; of mezianae Hagmann, Ilha Mexiana, 
mouth of Rio Amazonas, Brazil; of brasiliensis Liais, Rio Sado Fran- 
cisco, Minas Geraes, Brazil; of anulipes Hermann, neighborhood of 
Cuyabé, Mato Grosso, Brazil; of spegazzinit Ameghino, Rio Pescado, 
Departamento de Ordn, Salta, Argentina; of peruvianus Gray, ‘‘Peru- 
vian Amazons,” here restricted to Santa Cruz, Rio Huallaga, Pert 
(this locality that of a specimen reported by Bartlett (in Gray, Proc. 
Zool. Soc. London (1872), p. 625, 1872) as mother of type; of aenigma- 
ticus Gray, said to be ‘Sunia,” Cordillera Oriental, Ecuador, but 
almost certainly from Macas, eastern Ecuador; of ecuadorensis Gray, 
Macas, eastern Ecuador; of tapir Erxleben, suillus Blumenbach, and 
americanus Gmelin, here restricted to Surinam; of rufus Fischer, 
Guiana, probably French Guiana; of maypuri Roulin, here restricted 
to the Guianas; of guianae Allen, Tumatumari, British Guiana. 

Distribution —aAs for the subgenus except in western Colombia 
where it is replaced by another race (see map, fig. 61). Tschudi 
(supra cit.) affirmed that in Peri the common tapir is confined to the 


482 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Te 


Ficure 58.—Crania of American tapirs, lateral views X 4. a, Tapirus terrestris; b, Tapirus 
bairdit; c, Tapirus pinchaque. 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 483 


Ficure 59.—Crania of American tapirs, dorsal views X 4. a, Tapirus terrestris; b, Tapirus 
bairdii; c, Tapirus pinchaque. 


484 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


tropical forest region and is never found at 3,000 feet or more above 
sea level. On the other hand, Tate (supra cit., p. 223) stated that on 
“Mt. Auyan-tepui [Venezuela] tapir tracks were abundant as high as 
3,500 feet.” Tate added that “on the eastern slopes of the Andes 
near the Pastaza River I found tracks of tapir above 7,000 feet.” 
It is possible that the last refers to the mountain tapir of Ecuador, 
T. pinchaque. It has already been shown (p. 478) that the 
common tapir recorded from “Talamanca, Costa Rica,’”’ by authors, 
is probably South American, possibly Brazilian, in origin. 

Characters —Those of the subgenus; general external characters 
given under the generic heading. Skin of adult female from Surinam 
(USNM) blackish brown on back and sides, dark brown on chest, 
belly, and limbs; top of head blackish brown, ears blackish brown 
except for white edging; cheeks grizzled brown and gray, throat with 
more gray, neck brown, chin blackish brown; mane from front of ears 
to withers black. Specimen from Rio Yuruari, Venezuela, figured by 
Sclater as “ Tapirus dowii (?)’”, similarly colored but jaws strikingly 
white. Two skins of menagerie individuals (CNHM, received from 
Chicago Zoological Society) also like Surinam specimens, one practi- 
cally indistinguishable, the other with less black on sides and limbs. 
Untanned skin from Buena Vista, Santa Cruz, Bolivia (CNHM, 
collected by José Steinbach), considerably paler, dorsal surface from 
Cinnamon-Brown to Tawny, sides paler, mixed with gray, mane black- 
ish brown, limbs like back but becoming Prout’s Brown on posterior 
surface. Skin of adult male (USNM) that died at 32 years of age 
after living 27 years in the National Zoological Park, Washington, is 
extremely pale; back Cinnamon-Brown mixed with gray, sides more 
grizzled, underparts dominantly gray to dirty white, mane Prout’s 
Brown, hoofs unpigmented. A portion of hide from posteriormost 
end of body of another specimen (USNM) from same zoological park 
is nearly uniformly Ochraceous-Tawny on dorsal surface, sides 
grizzled, underparts tending to become dominantly gray; pelage of 
both menagerie specimens considerably thicker and longer than in any 
wild, living 7’. terrestris. 

Measurements.—Of an adult male and a female, respectively, col- 
lected in Porto Campo, Rio Sepotuba, Mato Grosso, Brazil, by Leo E. 
Miller (ex Allen, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 566, 1916): 
In millimeters, total length, 2,070, 2,000; tail, 60, 100; ‘‘hind foot” 
(digits only), 140, 140; ear, 120, 120. Of type of spegazzinw, collec- 
tor’s measurements cited by Ameghino (supra cit.): Height, 80 cm.; 
weight, approximately, 250 kilos. Of type of anulipes, living animal 
measured by Hermann (supra cit.): Total length, 210 cm.; height at 
shoulder, 77 cm.; height at rump, 83 cm. Of an adult from Pert, 
measurements from Tschudi (supra cit.): Head and body, 6 ft., 7 in. 


MAMMALS OF NORTHERN COLOMBIA—-HERSHKOVITZ 485 


(German system); tail, 3 in., 3 lines; height at withers, 3 ft., 6 in. 
Of an adult male from Paraguay (ex Azara, Essais sur |’Histoire 
Naturelle des Quadrupédes de la Province du Paraguay . . . , French 
ed., vol. 1, p. 5, 1801): Head and body, 2,000 mm.; tail, 100 mm.; 
height at shoulder, 1,100 mm., at rump, 1,130 mm.; circumference of 
chest, 1,215 mm.; ear, from crown, 80 mm. 

Remarks.—All evidence points to the existence of but one species 
of tapir east of the Andes in South America. That there is more than 
one subspecies within the range assigned to the typical form is likely 
but cannot be satisfactorily demonstrated without comparisons with 
the Linnaean terrestris from Pernambuco. Earlier characterizations 
based on single skulls only (laurillardi, spegazzinii, guianae, mexianae) 
define nothing more than individual variation. Descriptions of 
species based on skins of striped juvenals (aenigmaticus, ecuadorensis, 
peruvianus) and young adults with persistent juvenal striping 
(anulipes) are trivial. Except for the Guianan rufus, the remaining 
names included in the synonymy of terrestris are founded on biblio- 
graphic references or simply distaste for the original Linnaean 
designation. 

Color and, possibly, size seem to be the only valid characters for 
to ee subspecies of tapirs. Unfortunately, external charac- 
ters of typical representatives of terrestris are unknown. With their 
type localities now restricted, anta Zimmermann, sabatyra Liais, and 
laurillardi Gray become absolute synonyms of terrestris. The Rio 
Sao Francisco brasiliensis Liais is almost certainly identical with the 
Pernambuco form. Tapirs of the Guianas are extremely dark, 
blackish brown in general appearance, and probably distinctly darker 
than typical terrestris. The earliest available name for a Guianan 
tapir is tapir Erxleben (synonyms: suwillus Blumenbach, americanus 
Gmelin, rufus Fischer, maypuri Roulin, guianae Allen). It is ex- 
tremely doubtful if mexianae Hagmann, from the mouth of the Rio 
Amazonas, is recognizable. It may be referable to either the Guianan 
or typical form or it may represent an intergrading population not 
certainly separable from either of its nearest allies. In any case, the 
earlier named aenigmaticus Gray (ecuadorensis Gray) from Macas 
and peruvianus Gray from the Rio Huallaga must be given prior 
consideration if an Amazonian race is recognized. The tapir of Mato 
Grosso, Brazil, is extremely pale grayish brown in general appearance. 
Three adults collected by Colin C. Sanborn and mounted in a habitat 
group in the Chicago Natural History Museum agree with the pub- 
lished description of the living type of anulipes from Cuyabé, Mato 
Grosso. The specimen from Buena Vista, Santa Cruz, Bolivia, de- 
scribed above, tis similarly pale. This pale. austral tapir is We os 
distinguishable from the saturate Guianan form but comparison with 


486 PROCEEDINGS OF THE NATIONAL MUSEUM VOL 103 


the typical Pernambuco tapir is required. The name spegazzinii 
Ameghino (anulipes Hermann, a synonym) is available, should recog- 
nition be indicated. Finally, the tapir of the Maracaibo basin, 
western Venezuela, may be referable to the northern Colombian race 
but it is known from skulls only and is here provisionally assigned to 
the “catch-all” terrestris Linnaeus. 

Specimens examined.—Forty-nine. Braziu: Paré, 4 (CNHM); Rio 
Grandedo Sul, 4 (USNM); Mato Grosso, 1 (USNM);Descalvados, Mato 
Grosso, 4 (CNHM); 50 miles northwest of Miranda, Mato Grosso, 
1 (CNHM); Salto do Hua, Rio Maturaca, at Venezuelan boundary, 
Amazonas, 1 (USNM); Serra da Lua, Amazonas, 1 (CNHM); “Branch 
of Amazon,” 1 (USNM); Surram: Moengo, 1 (USNM); Paramaribo, 
1 (USNM); no precise locality, 2 (USNM). VEnrzurta: Empalado 
Savanas, Zulia, 1 (CNHM); Sierra de Perij4, Zulia, 1 (CNHM). 
Boutrvia: near Brazilian boundary, west of Sado Luiz de Caceres, 1 
(USNM); Buena Vista, Santa Cruz, 1 (CNHM). Pert: Tingo 
Maria, Huanuco, 1 (CNHM); Pozuzo, Huanuco, 10 (CNHM); 
Yarinacocha, Loreto, 1 (CNHM). ‘Costa Rica”: 1 (USNM, a skull 
only, probably from Brazil). Soura America: 11 (USNM, 9; 
CNHM, 2). 


Tapirus terrestris colombianus, new subspecies 
(“danta colombiana’’) 


Tapir [sp.] Roulin, Ann. Sci. Nat., Paris, ser. 2 (Zool.), vol. 18, p. 35, 1829 (dis- 
tribution of T. terrestris in Colombia: Lower Magdalena and Cauca valleys; 
common at Murindé, right bank of Rfo Atrato; in Colombian Darién but 
never seen on Panamanian side). 

Tapirus terrestris, Bangs, Proc. New England Zool. Club, vol. 1, p. 90, 1900 (adult 
female, skin and skull from Dibulla, coastal plain north of the Sierra Nevada 
de Santa Marta).—Allen, Bull. Amer. Mus. Nat. Hist., vol. 20, p. 480, 1904 
(two adults with skeletons, one spotted immature, from Cacagualito, Sierra 
Nevada de Santa Marta, 20 miles east of Santa Marta, altitude, 1,500 ft.).— 
Goodwin, Bull. Amer. Mus. Nat. Hist., vol. 87, p. 451, 1946 (Naranjo, Santa 
Marta region, Colombia). 

Tlapirus] terrestris, G. M. Allen, Extinct and vanishing mammals of the Western 
Hemisphere, p. 405, 1942 (part; Dibulla, northern Colombia.). 


Holotype-—Young adult male, skin and skull, USNM 281389; 
collected July 15, 1942, by Philip Hershkovitz; original number, 438. 

Type locality —E] Salado, eastern slope of Sierra Nevada de Santa 
Marta, on road between Valencia and Pueblo Bello, Department of 
Magdalena, Colombia; altitude, 430 meters. 

Distribution.—Tropical Zone of northern Colombia, in the Depart- 
ments of Magdalena, Atlaéntico (?), Bolivar, Cérdoba, and north- 
western Antioquia; the species is not known to occur west of the 


MAMMALS OF NORTHERN COLOMBIA—-HERSHKOVITZ 487 


Rio Atrato in the Department of Chocé and north of the Rio Ranch- 
erfa, in La Guajira. The subspecies may range into the Lake 
Maracaibo basin in Venezuela but external characters of the tapir 
there are unknown. (See map, fig. 62). H. H. Smith (in Allen, 
supra cit.) stated that on the western slope of the Santa Marta 
region, Magdalena, “the tapir is common from sea-coast to 6,000 
feet, and probably higher as I have seen tapir tracks at nearly 8,000 
feet.’”’ I saw no signs of tapir above 700 meters on the southern 
and eastern slopes of the Sierra Nevada de Santa Marta. In the 
absence of reliable records proving otherwise, the altitudinal range 
of the species has been determined as sea level to not over 1,200 
meters above. The tapir is certainly extinct now in the Department 
of Atléntico and is vanishing from Cérdoba. 

Characters.—Distinctly paler throughout than Guianan representa- 
tives of terrestris; slightly darker, less gray than the Mato Grosso— 
Chaco tapir at the opposite extreme of the range of the species. 


=> 


ca 
1, 


A 


Figure 60.—Structural variation in first upper premolar of American tapirs, X 2. a, first 
premolar of Tapirus terrestris with fully developed cinguloid shelf extending from proto- 
cone. 3, first premolar of Tapirus pinchaque, specimen from Ecuador (CNHM 47051) 
with cinguloid shelf absent (in another specimen from Colombia, CNHM 70557, premolar 
is as shown in a). 


Coloration of holotype-—Back thinly haired Prout’s Brown, basal 
portions of hairs buffy to ochraceous; side paler, hairs gray basally, 
Prout’s Brown to Cinnamon-Brown terminally; pelage of chest and 
belly thinner, more gray. Fore and hind limb Prout’s Brown with 
persistent juvenal spots and patches of whitish to buffy. Snout and 
frontal region Prout’s Brown, ear Prout’s Brown rimmed with white; 
cheek and side of lower jaw gray lightly mixed with brown, throat 
and anterior part of chest less brown; chin brown, lips fringed with 
stiff gray and buffy hairs. Sharply defined blackish brown mane 
extends from forehead to well behind middle of back. 

Measurements—Those of the holotype followed by those of an 
adult male paratype (in millimeters): Total length, 1,760, 1,870; 


488 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


tail, 46, 83; hind foot, 333, 350; ear, 137, 125; greatest length of 
skull (from gnathion to nuchal crest), 370 (m? functional), 385 (m3 
functional), zygomatic breadth, 175, 175. 

Remarks.—Distinction of colombianus from trans-Andean terrestris 
is based on comparisons with the blackish-brown representatives of 
the species in the Guianas (= Tapirus terrestris tapir ?Erxleben). 
The skin of the adult male paratype of colombianus could not be 
preserved and was discarded. Its color was quite like that of the 
type. The specimen recorded by Bangs (supra cit.) was collected 
by W. W. Brown, Jr., in Dibulla, a humid tropical locality on the 
northern base of the Sierra Nevada de Santa Marta. It was examined 
by Miss Barbara Lawrence of the Museum of Comparative Zoology, 
Harvard, and found to agree with the above description of colombi- 
anus except for its slightly paler back, chest, and belly, more buffy 
cheeks, sides of lower jaws, and throat; greatest length of skull 
(M? functional), 385 mm. 

Herbert H. Smith (in Allen, Bull. Amer. Mus. Nat. Hist., vol. 20, 
p. 431, 1904) reported that ‘‘all the hunters near Santa Marta aver 
that there is a tapir, found in the mountain forest, which, in general 
color, resembles JT. americanus, but has a broad white mark over the 
shoulder.””’ Smith concluded that it might represent an ‘‘undescribed 
tapir, which differs in color from all the known American species, 
and resembles that of the Malay Islands.’”’ The tapir in question, 
if not a myth, may be a pied individual of colombianus. 

Specimens examined.—Seven. Coutompsta: El Salado, Sierra Nevada 
de Santa Marta, Magdalena, 2 (USNM); El Orinoco, Rio Cesar, Mag- 
dalena, 2 (USNM); Rio Guaimaral, Rio Cesar, Magdalena, 1 (USNM); 
“Rio Magdalena,” 1 (USNM); Socorré, upper Rio Sint, Cérdoba, 
1 (CNHM). 

| Subgenus Tapirella Palmer 


Elasmognathus Gill, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, p. 183, 1865 
(genotype by monotypy, Elasmognathus bairdii Gill). 

Tapirella Palmer, Science, new ser., vol. 17, p. 873, 1903 (new name for Elasmog- 
nathus Gill, preoccupied by Elasmognathus Fieber, 1844, a genus of Hemip- 
tera). 

Included species —Tapirus bairdii. 

Distribution.—From México, in Veracruz and the Istmo de Tehuan- 
tepec, east into Campeche and British Honduras, south through 
Guatemala into Panamdé, Colombia west of the Rio Cauca, and Ecua- 
dor west of the Andes (see map, fig. 61). 

Characters.—Head flattened dorsally, a low mane, not always well 
defined, extending from front of ears to withers; proboscis longer and 
bulkier than in other American species; pelage of lowland populations 
thin and not completely hiding skin, in highland populations longer, 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 489 


thicker, and completely hiding skin; hoofs broad, larger than in sub- 
genus Tapirus with middle hoof always wider than long. Dorsal 
contour of skull (fig. 58, 6) flattened or slightly rounded; median 
frontal line usually placed abruptly above level of nasals, roughly 
parallel to horizontal plane of nasals but strongly divergent from hori- 
zontal plane of maxillary tooth row; superior longitudinal parietal 
ridges separated by a broad flat table, not uniting to form a sagittal 
crest (fig. 59, 5); ossification of vertical mesethmoid plate extending 
beyond tips of nasals to angle between premaxillae in old adults; outer 
anterodorsal surface of maxillae produced upward to form thin, 
parallel-sided plates embracing mesethmoid; posterolateral maxillary 
process projecting back to form inner lateral wall of narial meatus but 
not contacting nasal bone; posterolateral border of premaxilla rounded 
or angular and embraced by maxilla; nasal without descending process 
overlapping maxilla; two ossification centers of nasal sometimes per- 
sistent in fully ossified bone of adult. First upper premolar as in 
subgenus Tapirus; upper incisors orthodont. 

Remarks.—Tapirella resembles the Indian Tapirus (Acrocodia) 
indicus more than it does either of its American relatives. The raised 
but flat crown of Baird’s tapir has the same relationship to the equally. 
broad but low crown of the Indian tapir as the crested crown of 7. 
terrestris has to that of T. pinchaque. Cranial characters associated. 
with the proboscis are diagnostic of T. bairdit when compared with 
other American species but, in many details, are like conditions found 
in T. indicus. 

The name Tapirella, dimunitive of Tapirus, is most inappropriate 
for the largest living species of American tapir. 


Tapirus (Tapirella) bairdii Gill 
(Baird’s tapir; ‘“‘danta centroamericana’’) 


Tapir (Anta) Zimmermann, Geographische Geschichte . . . , vol. 2, p. 154, 1780 
(part; Yucatdin and Panamé4). 

Elasmognathus bairdii Gill, Proc. Acad. Nat. Sci. Philadelphia, vol. 17, p. 183, 
1865. 

Elasmognathus dowii Gill, Amer. Journ. Sci., ser. 2, vol. 50, p. 142, 1870. 

Elasmognathus bairdii, Hatcher, Amer. Journ. Sci., ser. 4, vol. 1, p. 175, pl. 3 
(nasals), pl. 4, fig. 4a, pl. 5, fig. 4, 1896 (osteological characters; phylogeny). 

Elasmognathus bairdi [sic], Gray, Proc. Zool. Soc. London (1867), p. 885, pl. 42, 
1868 (revision; description). 

Elasmognathus dowi {sic], Hatcher, Amer. Journ. Sci., ser. 4, vol. 1, p. 175, pl. 3 
(nasals), 1896 (osteological charaeters; phylogeny; ‘‘might better be con- 
sidered as a subspecies [of bairdiz]’”’). 

Tapirus (Elasmognathus) bairdi [sic], Sumichrast, Naturaleza (México), vol. 5, 
p. 332, 1882 (Sierra Madre, Istmo de Tehuantepec; Chiapas, México). 


490 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Tapirus bairdi [sic], Sclater, Proc. Zool. Soc. London (1872), p. 635, pl. 51, 1872.— 
Gaumer, Monografia de los mamfferos de Yucatdn, México, p. 43, 1917 
(Yucatan, México; British Honduras). 

Tapirus dow? [sic], Alston, Proc. Zool. Soc. London (1879), p. 666, 1880 (com- 
ments; range probably restricted to Pacific slope of Guatemala and Nic- 
aragua).—Alston, Mammalia, in Godman and Salvin, Biologia Centrali- 
Americana: Zoology, p. 104, pl. 8, figs. 3-5, 1882 (part; description, characters, 
bibliographic references; figure pl. 9= T. terrestris). 

Tapirus bairdii, Alston, Mammalia, in Godman and Salvin, Biologia Centrali- 
Americana: Zoology, p. 101, pl. 8, figs. 1, 2, 1882 (description, characters, 
bibliographic references).—Simpson, Bull. Amer. Mus. Nat. Hist., vol. 86, 
pp. 37-80, pl. 8, fig. 2, 1945 (osteological characters; comparisons; phylogeny). 

{Tapirella} dowi [sic], Simpson, Bull. Amer. Mus. Nat. Hist., vol. 86, p. 41, foot- 
note, 1945 (‘‘specific status does not appear to be demonstrated’’). 

Tapirella bairdii, Goldman, Smithsonian Misc. Coll., vol. 69, p. 81, 1920 (Pan- 
am4).—Goodwin, Journ. Mamm., vol. 27, p. 91, 1946 (dowii a synonym); 
Bull. Amer. Mus. Nat. Hist., vol. 87, p. 450, 1946 (Costa Rica: Talamanca 
and Pacuare, Limén; Carillo, San José).—Hatt, Cranbrook Inst. Sci. Bull. 33, 
p. 72, 1953 (México: Acttin Lara, Yucatdn; extinct). 


Lectotypes —Of bairdit Gill, skull only, adult, USNM 6019, collected 
April 9, 1863, by W. T. White (one of two cotypes designated by Poole 
and Schantz, U. S. Nat. Mus. Bull. 178, p. 233, 1942); of dowit Gill, 
skull only, young adult, USNM 11278, collected by J. M. Dow, 
original number, 1 (one of five cotypes designated by Poole and 
Schantz, loc. cit.) 

Type localities —Of bairdii Gill, “Isthmus of Panama,” here re- 
stricted to Canal Zone, Panama; of dowiz Gill, “Guatemala,” believed 
by Alston (Proc. Zool. Soc. London, 1879, p. 666, 1880) to be “‘confined 
to the Pacific slope of Guatemala and Nicaragua.” Later, Alston 
(Mammalia, in Godman and Salvin, Biologia Centrali-Americana: 
Zoology, p. 105, 1882) quoted Godman and Salvin as follows: ‘On the 
Pacific coast [of Guatemala] Tapirs are no doubt abundant. In the 
forest of the hacienda of El Overo, a few leagues from the port of San 
José, the proprietor, Don Juan Viteri, assured us that they were to be 
found in plenty. He it was, we believe, who supplied Captain Dow 
with the original specimens of 7. dowi [sic].’”’ Accordingly, the type 
locality of dowii is here restricted to the Pacific slope of Guatemala. 

Distribution —As for the subgenus. According to Hatt (Cranbrook 
Inst. Sci. Bull. 33, p. 72, 1953), who recorded skull fragments from 
caves in southern Yucatan, ‘‘tapir has not been known in Recent 
time from this part of Yucataén.’”’ In Panama the species is known to 
range from sea level to the summits of the highest ridges of the eastern 
and western mountains. Oliver Pearson, who climbed to the top of 
the Volcén de Chiriqui in 1937, found well-worn tapir trails at 3,350 
meters altitude, near the very summitofthe peak. (See map, fig. 62.) 

In northwestern Colombia, east of the Rio Atrato, Baird’s tapir 
lives side by side with Tapirus terrestris colombianus. The writer 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 491 


preserved a skull only of the Colombian tapir killed in the upper Rio 
Sint valley 3 kilometers below the mouth of the tributary Rio Verde, 
and a skull only of a Baird’s tapir killed by a hunter about 4 kilometers 
lower down the Sint, near the confluence of the Rio Nain. Within 
the same area, but more than a year before, a hunter killed a Baird’s 
tapir while it was feeding on fallen fruit of the cannon-ball tree 
(Couroupita guianensis) and two weeks later killed a Colombian tapir 
feeding on the same fruit. The hunter preserved the skulls and they 
were identified by the writer. 

Earliest authentic record of the existence of Baird’s tapir in South 
America and first knowledge of the occurrence of this species in western 
Ecuador is based on a photograph of the animal kindly loaned to the 
writer by Belle Benchley, Executive Secretary of the Zoological Society 
of San Diego. Mrs. Benchley added the information (in litt.) that the 
individual photographed was brought to the San Diego Zoological 
Park by Fred Lewis, who “had taken a small boat and gone up the 
river at Guayaquil, Ecuador, and brought it back.’”’ The animal died 
in captivity in 1945. 

Characters.—Those given for the subgenus. See also general exter- 
nal characters mentioned under the generic heading. Skin of adult 
male topotype of bairdii from Tiger Hill, C. Z., collected by E. A. 
Goldman, is dark brown with pelage thin, stiff, sleek, the skin showing 
through. Two skins of young adults from Guatemala, probably topo- 
typical of dowti, are darker, nearly black on dorsal surface, pelage 
thick, coarse, comparatively long, and completely hiding skin. 

Measurements—Of the topotype of bairdii from Tiger Hill: Total 
length, 2,000 mm.; tail, 70 mm.; hind foot, 372mm. Ofanadult from 
Vera Cruz, México (ex Goodwin, Bull. Amer. Mus. Nat. Hist., vol. 87, 
p. 450, 1946): Total length, 2,020 mm.; tail, 70 mm.; hind foot, 375 
mm.;ear,140mm. Skull lengths given under the generic heading. 

Remarks. —That bairdti and dowii are conspecific has already been 
indicated by Hatcher, Simpson, and, finally, by Goodwin, all cited 
above inthesynonymy. It is possible, however, that there may be two 
geographic races, one the typical lowland thinly haired form, the other 
a comparatively thickly haired highland race. However, additional 
characters to support what are ostensibly individual somatic responses 
to cooler climate are required for validating the name Tapirus bairdii 
dowii for the highland tapir of western Guatemala and El Salvador. 

The two Guatemalan skins described above were mounted specimens 
exhibited in the Columbian Exposition by the Guatemalan Commis- 
sion. They are now preserved as study skins in the collection of the 
U.S. National Museum and numbered 61221 and 61222, the last with 
skull. A skull only of a young individual, numbered 61221-B, has the 
same history and may correspond to the first skin. 


492 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 103 


The head of “ Tapirus dowi” figured by Alston (Mammalia, pl. 9, 
in Godman and Salvin, Biologia Centrali-Americana: Zoology, 1882) 
has the high, maned crest distinctive of T. terrestris, to which species it 
is now assigned. 

Baird’s tapir, the least known and the last living species of the genus 
to be given a Linnaean name, was the first recorded in European 
literature. A recognizable, though exaggerated, description was 
given by Peter Martyr D’Angher, the first chronicler of the discovery 
and conquest of America, in book 9 of the second (of eight) Ocean 
Decade of his ‘‘De Orbe Novo,” published in 1516. Martyr’s concept 
of the tapir, acquired from descriptions brought to him by the first 
explorers of the Isthmus of Panamé, is of an animal which “Nature 
created in prodigious form. It is as large as a bull, and has a trunk 
like an elephant; and yet it is not an elephant. Its hide is like a bull’s 
and yet it is not a bull. Its hoof resembles that of a horse, but it is 
not a horse. It has ears like an elephant’s though smaller and drooping, 
yet they are larger than those of any other animal.’’ Prior to official 
date of publication, the manuscripts of the ‘““Oceanic Decades” were 
made available to students and correspondents as they were being 
written, from 1494 onward. This led to a pirated published edition 
in 1504 and another in 1507, both Italian. The second record of a 
tapir refers to another Panamanian 7. bairdii and dates from the 
“Summario” of Gonzalo Fernéndez de Oviedo y Valdes, published in 
1526 (or 1525) at Madrid. The accurate description of this tapir is 
repeated, accompanied by a first-rate woodcut, in Oviedo’s “‘ Historia 
General y Natural de las Indias” (book 12, chap. 11, pl. 1, fig. 11), 
published in 1535 at Seville, Spain. Oviedo extolled the gastronomic 
virtues of tapir meat and slow-boiled tapir feet, and told of how the 
animal is hunted with dogs. During the remainder of the 16th 
century practically all travelers, missionaries, and students interested 
in New World natural history described or referred to Baird’s tapir. 
Francisco Hernéndez (Rerum Medicarum Novae Hispaniae The- 
saurus, seu Plantarum, Animalium .. ., tract. I, cap. 8, p. 3, 1651) 
described the Tlacazolotl, which, in spite of the long tail attributed 
to it, is unmistakably a tapir. The animal was said to occur in 
Atzcén [Veracruz], Tepotzotlén [México] and Tlaquilapaén [Hidalgo 
or Veracruz]. The Herndndez account of the Tlacazolotl is iden- 
tical to that of Fray Bernardino de Sahagtin in his “Historia 
General de las Cosas de Nueva Espana.’”’ This work was written 
during the latter half of the 16th century. The manuscript, ex- 
amined and cited by 16th and early 17th century students of 
Mexican history, was not published until 1831. Publication dates 
are of no importance in comparing the works of Hernandez and 


MAMMALS OF NORTHERN COLOMBIA—HERSHKOVITZ 493 


Sahagtn because they were written contemporaneously. It cannot 
be determined now which author, or editor, copied from the other. 

The first Linnaean reference to Baird’s tapir is by Zimmermann in 
1780 (supra cit.), who considered it the same as the Brazilian species. 
Zimmermann’s Panama record for the tapir is based either on Oviedo 
or other pre-Linnaean authors who quoted, Oviedo. His Yucatan 
record is almost certainly derived from the account of the tapir in 
“‘Dampier’s Voyages,” volume 2, part 2 entitled ‘“‘Two Voyages to 


=. Topirus bairdir 
= Jopirus terrestris colombianus 


Ficure 61.—Distribution map of American tapirs. Range of Tapirus terrestris terrestris as 
outlined includes those of other possibly recognizable subspecies. Type localities: (1) Per- 
nambuco, Brazil, Tapirus terrestris terrestris Linnaeus; (2) Surinam, [T. terrestris] tapir 
Erxleben; (3) Macas, Ecuador, [T. terrestris] aenigmaticus Gray; (4) Rio Pescado, 
Salta, Argentina, [T. terrestris] spegazzinii. 


494 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 103 


Campeachy; with a description of the Coasts, Product, Inhabitants, 
Logwood-Cutting, Trade, Etc. of Yucatan, Campeachy, New Spain, 
Ete.,’”’ page 102, which was first published in 1698. 

The common name “danta” for the tapir is a corruption of the 
Spanish word for elk, ‘‘alce” or “‘anta.”” Early Spaniards in America 
gave peninsular names to all animals that resembled, whether in fact 
or fancy, those they had known in the Old World. The now univer- 
sally accepted term ‘‘tapir” is from the same word in the Brazilian 
Tupi language. 


EXPLANATION OF MAP, FIGURE 62 
@ =Tapirus bairdii Gill 
Locality records of authentically identified specimens from South America, Panam4, and 
Costa Rica. Range of species extends northward into México. 
Costa Rica 
1. Pacuare, Limon. 
2. Talamanca, Lim6n. 
3. Carillo (=Carrillo), San José. 
PanaMA 
4. Volcan de Chiriqui, Chiriqui. 
5. Canal Zone (type locality). 
6. Mt. Pirri and Cana, Panama. 
CoLomBIA 
7. Unguia, Chocé (sight record, by author), 
8. Upper Rio Sint, Cordoba. 
Ecuapor 
9. North of Guayaquil (exact locality unknown). 
O=Tapirus terrestris colombianus, new subspecies. 
CoLoMBIA 
1. Dibulla, Magdalena. 
2. Cacagualito, Sierra Nevada de Santa Marta, Magdalena. 
3. El Salado, Sierra Nevada de Santa Marta, Magdalena (type locality). 
4. El Orinoco, Rio Cesar, and Rio Guaimaral, Magdalena. 
5. Socorré, upper Rio Sina, Cérdoba. 
6. Murind6é, Chocé. Recorded by Roulin as T. terrestris but identification 
questionable. 
Wi = Tapirus pinchaque Roulin. 
CoLoMBIA 
1. Santander (Department), Cordillera Oriental, at Venezuelan boundary. 
2. Las Juntas, upper Rio Combeima, southern foot of Mt. Tolima, Cordillera 
Central, Caldas. 
3. Paramo de Sumapaz, Cordillera Oriental, Cundinamarca (type locality). 
4, Volcan de Puracé, Cordillera Central, Cauca. 
5. Rio Majuas, upper Rio Magdalena, Cordillera Central, Huila. 
Ecuapor 
6. Borja, about 60 kilometers east of Quito, Cordillera Oriental Napo-Pastaza. 
7. Papallacta, Cordillera Oriental, Pichincha. 
8. Cordillera de los Llanganates, Cordillera Oriental, Tungurahua. 
9. Paramo de Azuay, Cordillera Oriental, Azuay. 


MAMMALS OF NORTHERN COLOMBIA—-HERSHKOVITZ 495 


Specumens examined.—Sixty. CoxtompBia: Unguia, Urabé, Chocé, 
1 (CNHM). Panam&: No precise locality, probably Canal Zone, 
5, including type of bairdit (USNM); Tiger Hill, C. Z., 1 (USNM); 
Madden Dam, C. Z., 1 (USNM); Rio Chagres, C. Z., 1 (USNM); 
Panama Railroad, C. Z., 1 (USNM); Gattin, C. Z., 1 (USNM); Mt. 
Hope, C. Z., 1 (USNM); Mt. Pirri, Panamé, 1 (USNM); Cand, 
Panama, 2 (USNM). Costa Rica: Talamanca, 4 (USNM); Pacuare, 
3 (USNM); San José, 1 (USNM); “Dota Mountains,” 1 (USNM); 
no precise locality, 6 (USNM). Nicaracua: Ebenezar, Rio Prinza- 


Tapirus bairdii 
Tapirus terrestris colombionus 
Tapirus pinchaque 


Figure 62.—Map of locality records for tapirs in Colombia and adjacent countries; type 
localities circled. See opposite page for key. 


496 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 103 


polka, 1 (USNM). Honpuras: No precise locality, 2 (USNM). 
Ex Satvapor: No precise locality, 4 (USNM). Guatrmata: Los 
Amates, Yzabal, 2 (CNHM); no precise locality, 8 (USNM). 
British Honpuras: Belize, 1 (CNHM); Middlesex, 2 (CNHM). 
México: Achotal, Vera Cruz, 2 (CNHM); Istmo de Tehuantepec, 
1 (USNM); Chiapas, 1 (USNM); Buena Vista, 1 (USNM). CrntTrAu 
America: No precise localities, 5 (USNM). 


®, S, GOVERNMENT PRINTING OFFICE: (984 


PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


issued { 
we'd 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1954 No. 3330 


A REVISION OF THE GOATFISH GENUS UPENEUS WITH 
DESCRIPTIONS OF TWO NEW SPECIES 


By Ernest A. Lachner 


The most recent and comprehensive studies of the genus Upeneus 
were included in the faunistic reports of Herre and Montalban, 1928, 
Weber and de Beaufort, 1931, and Fowler, 1933. Six species of 
Upeneus were recognized by Herre and Montalban as occurring in 
the Philippine Islands, and Weber and de Beaufort recognized the 
same species in the Indo-Australian Archipelago. Fowler listed eleven 
species in his Philippine report but included five extralimital species, 
four of which are not valid or are highly questionable. 

In all, 24 nominal species have been referred to the genus and much 
nomenclatorial confusion exists. The characters presented by the 
various authors to distinguish the species did not prove satisfactory 
in the identification of specimens in the U. S. National Museum from 
the same faunal areas. Additional specimens from the Philippine 
Islands and the Persian Gulf did not conform to any published 
accounts. 

The object of this study is to determine the valid species in the 
genus and characters for their accurate identification, to evaluate the 
extent of interspecific differentiation, and to evaluate the population 
divergence in the various subfaunal areas. 

Ten species are herein recognized, two of which are described as new. 

The collections in the U. S. National Museum formed the chief 
basis of this study on which counts, measurements, and color analyses 
were made. These collections are listed by subfaunal areas of the 
Indo-Pacific region in the descriptions of the respective species. 

279126—54——1 497 


498 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Specimens and types at the Museum of Comparative Zoology, Harvard 
University, were also examined. One specimen (Upeneus subvittatus 
Snyder, 1907, p. 101) was loaned from the Natural History Museum, 
Stanford University, through the courtesy of Dr. George S. Myers. 
The type of Mullus dubius Temminck and Schlegel (1843, p. 30) was 
reexamined for the author by Dr. M. Boeseman, Rijksmuseum van 
Natuurlijke Historie, Leiden, Netherlands. 

T am also indebted to William C. Schroeder for making laboratory 
facilities available at the Museum of Comparative Zoology, Harvard 
University, and to Dr. Robert R. Harry, Academy of Natural Sciences, 
Philadelphia, for loan of the holotype of Upeneus phillipsi Fowler. 
The photographs were taken by personnel of the Smithsonian Insti- 
tution’s photographic laboratory. 

The synonomy listed in the descriptive accounts of each species is 
incomplete. Many faunal studies were largely ignored except in cases 
where adequate descriptions or illustrations were presented, or where 
the collections involved were available for reexamination. The geo- 
graphical distribution given for each species was constructed from the 
distributions of examined specimens and from the literature where 
the descriptions or illustrations were sufficiently thorough and accurate 
to enable me to identify the species with assurance. 

All measurements of the length of a fish refer to the standard length, 
unless stated otherwise. The method of counting fin rays, gillrakers, 
and scales of the body is similar to that given by Lachner (1951, p. 
581). In the first paragraph under “Description” of each species the 
count for each character is recorded as follows: the mean: range 
(number of specimens) ; for example, pectoral rays 16.1: 15 to 17 (68). 
This method of recording does not apply for the new species. For 
these, separate methods are given in the description of each species. 
Dark spots at the tips of the lobes of the caudal fin, distinct from 
black margins, were counted as bars in the tabulated data, but the 
small spots or blotches near the midbase of the caudal fin were 
omitted. 


Diagnostic characters 


Several of the important diagnostic characters useful in distinguish- 
ing the species have been misinterpreted by various authors or were 
entirely overlooked. It is necessary, therefore, to discuss the critical 
characters in the approximate sequence of their importance in the 
identification of the species. These characters are of taxonomic value 
because of their low variability and the accuracy by which they can 
be measured. The sequence in which they are discussed is not 
intended to portray phylogenetic relationships although this may 
partially exist. 


GOATFISH GENUS UPENEUS—LACHNER 499 


The number of SPINES IN THE SPINOUS DORSAL FIN is either 7 or 8 
in any particular species. Only one abnormally developed specimen 
of a species normally with seven had 6 spines. The difference of one 
spine among the species is associated with the presence or absence 
of the minute first spine. No variability was found in this character 
in any species in more than 300 specimens examined. This small 
spine is located near the first enlarged one, often partially embedded, 
and may be entirely overlooked without employing some probing 
and a microscope. The variability of this character as given by 
Fowler (1933, pp. 322, 341) is erroneous. 

Counts of the PECTORAL FIN RAYS for 10 species from various locali- 
ties in the Indo-Pacific are given in table 1. The range of this charac- 
ter in any species did not exceed four rays. An inspection of the data 
in the table shows that the species are divisible into two groups and 
that the modes of the frequency distributions fall on 13 or 14 in one 
group and on 16 in the other, except for U. parvus, where only 6 speci- 
mens were available for study. It is unfortunate that descriptive 
accounts of certain nominal species did not include this character; 
with it, a more reliable interpretation would have been possible. 

The TOTAL NUMBER OF GILLRAKERS of 260 specimens are arranged 
from lowest to highest, respectively, for nine species from the Indo- 
Pacific region in table 1. The tenth species, parvus, represents the 
single form in American waters. These data represent specimens from 
various localities. 'The range of this character for each species is low. 

The usual count of the gillrakers reported in the literature is that 
of the lower limb only. The rudiments were seldom included. Some- 
times they were partly included but without explanation as to how 
they were distinguished from the developed rakers, making it impos- 
sible to interpret the count. To form a more reliable basis for the 
interpretation of data in the literature, the raker count of both limbs 
and the raker-rudiment relationship are recorded separately in tables 
2,4, and 5. It is apparent from these data that the number of rudi- 
ments and rakers differs for different species and that the addition of 
developed rakers and rudiments of the upper and lower limbs to form 
the total count displays the greatest differences among the species. 

Species having a minute first dorsal spine can be divided into two 
subgroups on the basis of the total number of gillrakers (see key, 
p. 508). The number of gillrakers in these two divisions also has 
a positive relationship to the number of pectoral fin rays. 

The species are divisible into two groups on the basis of color of 
peritoneum: Those in which it is light to silvery (first four species 
and the last species, table 1), and those in which it is light brown to 
blackish (all other species, table 1). A positive relationship occurs 


103 


VOL. 


PROCEEDINGS OF THE NATIONAL MUSEUM 


500 


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GOATFISH GENUS UPENEUS—LACHNER 501 


between color of peritoneum and number of gillrakers (table 1) for 
the Indo-Pacific species but does not hold when the Atlantic species, 
U. parvus, is included. Indo-Pacific species with the peritoneum 
light colored have fewer gillrakers than those with dark peritoneal 
linings. 

The COLOR OF THE PERITONEUM is a clear-cut character in most of 
the species and specimens. However, certain species show some varia- 
tion and intermediacy and these may be difficult to evaluate, espe- 
cially to one inexperienced with this character. Species with light or 
transparent to silvery peritoneal linmgs may have a few scattered, 
dark, pepperlike spots. Species with peritoneum colored light brown 
to blackish may in some cases have a silvery cast over a brownish 
background. This is especially so in asymmetricus and in some 
small specimens of bensasv. 

A summary of the number of ROWS OF VERTICAL SCALES in 253 
specimens and scales below the lateral line is given in table 2. The 
low intraspecific variability of the number of scale rows is evident in 
that the range does not exceed five for any species. The scales are 
somewhat deciduous and the count cannot always be made. Many 
large or poorly preserved specimens lack some or all of the scales. 
The number of scale rows has been widely used in the literature as a 
diagnostic character but interspecific differences have never been com- 
pared. The discrepancies between my data and that given in the 
literature may in part be associated with different methods of making 
this count. Others are not explainable. 

Life colors are useful in the identification of the species but fade in 
preservation. However, certain brown to black stripes, bars, spots, 
and blotches on the body and fins persist in preservation in varying 
intensities. These COLOR MARKS are extremely valuable in distin- 
guishing certain species, especially the number of oblique bars on the 
upper lobe, lower lobe, or both lobes of the caudal fin. In some spe- 
cies the number of bars increases with growth, whereas in others it 
remains constant. The variation and relationship of the number of 
bars in respect to size of body for three species is given in table 3. 
Failure to understand this character by various authors has caused 
considerable confusion in distinguishing the species (Fowler, 1918a, 
Devs, -hig..15). 

Average values of the LENGTH OF THE BARBEL show differences 
among the species, but great overlaps occur in the ranges (table 1). 
In at least one species (swlphureus) there may be an increase in the 
length of barbel with increase in length of body (see tables 6, 8, 9, 11). 

Other characters such as depth of body, length of head, length of 
snout, and size of eye were investigated. Although small average 
differences were apparent among the species, they were too insignif- 


502 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


' 
Taste 2.—Number of vertical scale rows and scale rows below the lateral line, and © 
number of gillrakers on each limb of the first arch in 10 species of Upeneus | 


Scales below 


Vertical scale rows laters’ tne | 


Species 


28 | 29 | 30 | 31 | 32 | 33 | 34 | 35 | 36 | 37 | 38 6 fi 


asymmetricus 
INOULOCEN SE ee Apt a ee ee aor ale el erie ee eee 
ORDER ek cre re | ee ee eee | nee eres 
sulphurews 45+ SPe8e sree th oo ree ese Sete ||Ss oF beeen lee 
DOTOUS Scant eee Soe ee or een eee neee Ren eee ea ne aaa eee | aes | eae 
Gillrakers 
Species Upper limb ! Lower limb ! 
| 

5| 6] 7 8 9 | 10 13 14] 15 16 | 17 18 | 19 | 20/21 22 
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moliccensisu tis TS aera eee a Pe hl ie ts Ya ees | a | pS |e I} Sits sa 2 2 
Oitaris LEVEL Ta Shy PATS ST EIA ee: BO CbSr| Wate ees) See eae eos 1 | 14):34) 17) 1 1 
SULDNUT CUS ons sence e ane eee eee Eee Za 20) AS Efe e ce tole eae 3) 8 |-20] 10 2 
paronsy_ LUST eee if tN RS epee WS EEE RS ee eS ee ee eee || oe 7 Di) 4 peLGs pss See ee 


1 Raker at angle of arch omitted from counts, all rudiments included. 


TABLE 3.—Variation and relationship of the number of bars on the caudal fin with 
increase in body length in three species of Upeneus 


Number of caudal bars on upper lobe—on lower lobe: ! 
Length in mm. 


1 Numbers in parentheses refer to U. oligospilus; those without enclosures, U. ¢tragula; and those in brac- 
kets, U.luzonius. Following data not recorded for U. tragula: 1 specimen, 97 mm., 4 bars on upper lobe, 
6 on lower lobe; 1 specimen, 107 mm., 3 bars on upper lobe, 5 on lower lobe. 


GOATFISH GENUS UPENEUS—LACHNER 503 


icant or variable to be of any practical taxonomic importance. The 
basal rigidity of the barbel, used as a key character by Weber and de 
Beaufort (1931, p. 363), was not found reliable. The different lengths 
of the barbel among the species probably led them to misinterpret 
this character. Herre and Montalban (1928, pp. 96-97) distin- 
guished between groups of species by the presence or absence of pre- 
orbital scales. While this character may be of some value, great 
errors in the identification of the species can result since these scales 
as well as those on the head and body are somewhat deciduous in 
most species. The size and number of intestinal loops and the num- 
ber of pyloric caeca were studied but poor preservation of these 
structures made it impossible to evaluate their taxonomic significance. 


Group relationships 


The species are divisible into two groups in three different ways 
by using a single character or combinations of characters. The first 
method, based chiefly on the absence of the minute first dorsal spine, 
separates bensasi, asymmetricus, and parvus from the remainder of 
the species, in which the spine is always present. ‘These three species 
have little or no coloration on the dorsal fins (limited to light tan 
spots arranged in rows in asymmetricus and very faint bars in parvus), 
whereas the alternate group has the dorsal marked with bars or 
blotches, always very distinct except in arge and luzonius. 

A second method includes the combination of the color of the peri- 
toneum and the number of gillrakers. The Indo-Pacific species 
luzonwus, arge, oligospilus, and tragula are related in having, almost 
always, light to silvery colored peritoneal linings, and fewer and 
shorter gillrakers. The remainder of the Indo-Pacific species have a 
light brown to blackish peritoneum, a higher number of gillrakers, 
and comparatively longer rakers. Of these species, bensasi is inter- 
mediate in respect to the number of gillrakers and bensasi and asym- 
metricus show the greatest variation in the color of the peritoneum. 
The Western Atlantic form, parvus, does not conform to this rela- 
tionship in that the peritoneum is light but the gillrakers are 
numerous. 

The third method, based on the number of pectoral fin rays (table 
1), clearly indicates two groups of species. 

Although the three methods are useful in identifying the species, 
the presence or absence of the minute first dorsal spine strongly sug- 
gests an ontogenetic change resulting in two phyletic lines on a sub- 
generic level of organization. I do not propose or advocate sub- 
generic terminology in a problem such as this where so few species 
are involved and where troublesome and complicated classification 
problems above the species level do not exist. 


VOL. 108 


PROCEEDINGS OF THE NATIONAL MUSEUM 


504 


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505 


GOATFISH GENUS UPENEUS—LACHNER 


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279126—_54——_2 


506 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Populations 


No notable differentiation of any species in the subfaunal areas of 
the Indo-Pacific was revealed by an analysis of various meristic counts 
(see descriptions of bensasi, vittatus, and tragula, and tables 6, 10, and 
11), proportional measurements, color, and color pattern of specimens 
segregated by locality. Differentiation in this genus appears to have 
gone to the species level, after which considerable stability was at- 
tained. These conclusions are tentative owing to the limited number 
of specimens of several species. 

Although little is known of the life histories of these inshore forms, 
populations of such widely distributed species as vittatus, sulphureus, 
and tragula occurring in the subfaunal areas of East Africa, the East 
Indies, the Philippines, and areas of Oceania are assumed to be 
considerably isolated, yet they are characteristically homogeneous. 

Larval forms of some species are thought to be pelagic, and their 
drift or movement may account for some minor association of these 
populations. 

The genus is restricted to the tropical and subtropical, littoral, 
marine waters of the Indo-Pacific and Western Atlantic regions. The 
East Indies and Philippine Islands, near the center of the Indo-Pacific 
region, contain at least seven of the species, four are known from 
East Africa, four from Oceania, but only one (arge) is known in the 
Hawaiian fauna. Five extend northward as far as southern Japan 
and at least three reach eastern Australia. Only one species, U. 
parvus, occurs in the Western Atlantic area of American waters. 


Genus Upeneus 


Upeneus Cuvier and Valenciennes, 1829, p. 448 (type species, Mullus vittatus 
Forsk&l, designated by Bleeker, 1876, p. 333). 

Hypeneus Agassiz, 1846, p. 190 (type species Mullus vittatus Forsk&l) (cor- 
rected orthography). 

Upeneoides Bleeker, 1849, p. 64 (type species, Mullus vittatus Forskal, designated 
by Jordan, 1919, p. 240). 

The characters best defining the genus are: (1) dentition complete, 
consisting of small villiform teeth on the vomer in the form of an 
irregular or triangular patch, on the palatines in an elongate band, 
and on both jaws in bands of narrow to moderate widths, and (2) 
scales present on soft dorsal, anal, and caudal fins. The bodies are 
elongate and somewhat compressed ; the caudal fin is marked with dark, 
oblique bars in most species. The species attain a small size compared 
with other members of the family. 

The following characters, some of which apply to other genera in 
the family, were found to be common to all the species: Anal fin rays 
I, i, 6: pelvic fin rays I, 5; caudal fin rays i, 7-++6, 1; scale rows around 


GOATFISH GENUS UPENEUS—LACHNER 507 


caudal peduncle 16; scales ctenoid with 4 to 7 radii; lateral line 
complete; first elongate spine of first dorsal fin flexible; a small portion 
of barbel, less than one-third its total length, rigid at base. 


Questionable species 


I place two nominal species in a doubtful status, Upeneoides sun- 
daicus Bleeker (1855, p. 411; 1877, pl. (4) 394, fig. 2) and Upeneus 
taeniopterus Cuvier and Valenciennes (1829, p. 451, type locality 
Trinquemale, Ceylon). 

It is highly possible that U. sundaicus Bleeker may be represented 
by either U. tragula or luzonius. Bleeker’s specimens were taken in 
the East Indies where both tragula and luzonius occur. Weber and 
de Beaufort (1931, pp. 370-371) saw one of Bleeker’s specimens but 
their account, at least in part, was extracted from Bleeker. The 
account of sundaicus by Herre and Montalban (1928, p. 98) was 
“compiled from Bleeker, and Evermann and Seale” (1907, p. 88). 
Fowler’s account (1933, p. 323) was also compiled from Bleeker. 
Examination of the specimen reported by Evermann and Seale as 
sundaicus (orig. No. 3201, USNM 56138) revealed a large, very poorly 
preserved specimen of luzonius. U. luzonius usually has weakly 
developed bars on the upper caudal lobe while those on the lower lobe 
may be completely obscure. Three dark saddles on the body are 
often completely faded. Such specimens, as well as large faded 
specimens of tragula, could easily have been involved in Bleeker’s 
illustration. Yet, there still is the possibility of the existence of a 
species unknown to us. There is no method of solving this problem 
at present. Intensive collecting in the East Indies Islands, as well as 
study, is necessary. The characters listed for sundaicus by Bleeker 
and Weber and de Beaufort (dorsal spines VIII, the first spine minute, 
“oillrakers 13-+3,” barbels reaching “hindborder of preoperculum’’) 
clearly relate it with tragula and luzonius. The number of vertical 
scale rows that they report, 33 to 35, is higher than in these species, 
but this may be due to the different methods of counting. Only the 
bars on the caudal fin illustrated by Bleeker are unique but these 
may be highly diagrammatic. Therefore, sundaicus may be a syno- 
nym of tragula, may replace luzonius, or may represent a distinct 
species. 

The second questionable form, Upeneus taeniopterus, known only 
by the type specimen, is characterized by having seven dorsal spines, 
each dorsal fin with three bars, each lobe of the caudal fin with six 
oblique bars, and a large triangular reddish spot on the caudal fin. 
Day (1876, p. 122) reported on the faded type and Fowler (1928, p. 
227; 1933, p. 327) repeated Day. Steindachner (1901, p. 487) re- 
ported two specimens from Honolulu, but he probably had U. arge, 
the only member of the genus found in the Hawaiian Islands to date. 


508 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Key to the species of Upeneus 


(References to groups of species in the descriptions pertain to all the species in the following categories of 
the key: 2a, bensasi group; 4a, vittatus group; 4b, tragula group.) 


la. Dorsal spines VII, the first spine longest. 
2a. Pectoral rays number 13 or 14; vertical scale rows range from 28 to 31; 
peritoneum brown to silvery brown. 
3a. Lower lobe of caudal fin without oblique bars, upper lobe with 2 to 3 
faint, oblique brownish or dusky bars; body nearly uniform light tan 
to brown, without lateral stripe or dark brown saddle posterior to 
soft dorsal fin; fewer gillrakers, modally 25, range from 23 to 27. 
U. bensasi (Schlegel) 
3b. Lower lobe of caudal fin with 6 or 7 narrow, brown, oblique bars, and 
the upper lobe with 3; body dark tan above, light tan below with 
a brown, horizontal stripe on midside and a dark brown saddle on 
caudal peduncle just posterior to soft dorsal fin; more gillrakers, 
modally 28, range 27 to 29____-_-_- U. asymmetricus, new species 
2b. Pectoral rays number 15 or 16; vertical scale rows range from 36 to 38; 
peritoneum light to silvery; upper and lower lobes of caudal fin with 3 
oblique, brownish black bars on the smaller specimens (68, 69 mm.) 
increasing to 5 bars on the larger specimens; bars more pronounced on 
lower lobe; body light tan, possibly with light colored median stripe; 
without dark saddle over caudal peduncle; gillrakers range from 26 to 29. 
U. parvus Poey 
1b. Dorsal spines VIII, the first spine minute. 
4a. Total number of gillrakers range from 26 to 32; pectoral rays number 15 
to 18; peritoneum brown to black. 
5a. Caudal fin transparent to dusky, without dark bars; chin barbels long, 
58 to 82 percent of head length in large specimens (over 85 mm.); 
barbel when extended posteriorly usually extends beyond vertical 
drawn through posteriormost point of preopercle. 
U. sulphureus Cuvier and Valenciennes 
5b. Caudal fin with oblique dark bars on upper lobe; chin barbels short, 
46 to 66 percent of head length; barbel when extended posteriorly not 
reaching vertical drawn through posteriormost point of preopercle. 
6a. Lower lobe of caudal fin transparent to dusky, without dark, oblique 
bars; a pale to yellow, median horizontal stripe on side of body, often 
faintly developed, or obscure___-_----- U. moluccensis (Bleeker) 
6b. Lower lobe of caudal fin with 2 to 3 dark oblique bars, the outer bar 
widest and more intensely colored; a light colored median and 
dorsolateral, horizontal stripe on body usually present. 
U. vittatus (Forsk4l) 
4b. Total number of gillrakers range from 19 to 25; pectoral rays number 12 to 
15; peritoneum silvery to transparent, sometimes with scattered, fine, 
brownish spots. 
7a. Scales small, 36 to 38 vertical rows on body, 7 rows below lateral line; a faint, 
tan colored median and dorsolateral, horizontal stripe on body, often 
completely faded_...._-.....---- U. arge Jordan and Evermann 
7b. Scales large, 28 to 32 vertical rows on body, 6 rows below lateral line; 
a conspicuous dark brown median stripe on body always present. 
8a. A dark brown saddle just posterior to base of soft dorsal fin almost 
always present; two additional saddles through spinous and soft 
dorsal fins sometimes evident; spotting or blotches absent on body; 


GOATFISH GENUS UPENEUS—LACHNER 509 


upper lobe of caudal fin with 6 brown, oblique bars in adult specimens 
of about 80 mm. in length, 4 to 5 bars in juveniles of about 50 mm.; 
lower lobe of caudal with 6 to 7 such bars in adults, 4 to 6 in juveniles; 
second spine of spinous dorsal fin usually longest; dark spots or 
blotches on spinous dorsal fin almost always faded; barbels long, 
usually reach vertical drawn through most posterior portion of 
preopercular margin; barbel length in percent of head length ranges 
PrOMMGE $0 Was oe eh ye es U. luzonius Jordan and Seale 
8b. Dark brown saddle just posterior to base of soft dorsal fin faint, 
usually not visible; saddles through fins absent; numerous, distinct, 
small brown spots on cheeks and sides of body to belly; upper lobe 
of caudal fin almost always with 4 to 5 brown, oblique bars in adults 
over 80 mm. in length, 3 or 4 in juveniles less than 80 mm. (see table 
3); lower lobe with 5 or 6 brown, oblique bars in adults, 4 or 5 in 
juveniles; fourth spine of spinous dorsal fin longest or about equal to 
third; dark spots or blotches on spinous dorsal fin almost always 
conspicuous; barbels short, not reaching vertical drawn through 
most posterior portion of preopercular margin; barbel length in 
percent of head length ranges from 52 to 68__U. tragula Richardson 
8c. Dark brown saddle just posterior to base of soft dorsal fin faint or 
completely obscure; saddles through fins absent; a few small dusky 
spots on sides of head, almost completely absent on sides of body to 
belly but with large, irregular, dusky to blackish blotches on body; 
upper and lower lobes of caudal fin with 3 or 4 dusky to black, 
oblique bars in adults over 80 mm. in length, 3 in juveniles; third 
spine of spinous dorsal fin longest or about equal to fourth spine; dark 
blotches on spinous dorsal fin almost always conspicuous; barbels 
short, not reaching vertical drawn through most posterior portion 
of preopercular margin; barbel length in percent of head length ranges 
from; 50 4h0, 642023 e2ae oes ee on U. oligospilus, new species 


Upeneus bensasi (Temminck and Schlegel) 
PuatE 13, Figure A 


Mullus bensasi Temminck and Schlegel, 1843, pt. 2, p. 30, pl. 11, fig. 2 (type 
locality, Nagasaki).—Boeseman, 1947, p. 43. 

Upeneoides gutitatus Day, 1867, p. 938, (type locality, Madras?); 1876, p. 121. 

Upeneoides japonicus Steindachner and Déderlein, 1884, p. 22 (type locality, 
Tokyo, Kochi, and Tango, Japan) (not Mullus japonicus Houttuyn, 1782, p. 
334). 

eee tokisensis Steindachner and Déderlein, 1884, p. 22 (name in synonymy, 
specimens from Tokyo). 

Specimens studied—One hundred fifteen specimens, ranging in 
length from 37 to 148 mm., from the following localities: Japanese 
area, 16 USNM collections, 42 specimens; Formosa, 3 USNM collec- 
tions, 21 specimens; Philippine Islands, 8 USNM collections, 52 
specimens. 

Description.—Dorsal rays, VII-i,8(65), the first spine longest 
(two abnormal specimens with some irregularly developed spines, 
VIII-i,8 and VI-i,8, the first spine in both specimens also longest; 
pectoral rays 13.8: 13 to 14 (44); vertical scale rows 29.4: 29 to 31 


510 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


(25); scale rows above lateral line 3 (7); scale rows below lateral line 
6.1: 6 to 7 (15); total number of gillrakers 25.0: 23 to 27 (38); length 
of longest raker in longest filament about 1.1 to 1.3 (6). 

Peritoneum brown to blackish; preorbital scales present; barbels 
long, extending beyond preopercular margin, barbel length in percent 
of head length 64 to 84 (35); first dorsal spine longest. 

Color in alcohol.—Head and body uniform tan to brown above and 
light tan below; some dusky pigmentation on outer portion of soft 
dorsal fin in larger specimens and two faint, dusky, horizontal bars 
sometimes seen in smaller ones, 40 to 60 mm. in length; lower lobe of 
caudal fin dusky, the tip of the rays transparent to light tan, the 
smaller specimens have a lengthwise dusky streak through middle 
portion of lobe; 3 faint oblique brownish to dusky bars on upper lobe 
in larger specimens and 2 bars in smaller ones about 40 mm. in length; 
remainder of fins transparent to uniform light tan. 

Geographical distribution.—This species has been reported from the 
East African coast eastward through the East Indies and Philippines, 
and northward to Formosa, east China, Ryukyu Islands, and southern 
Japan. It has not been reported from the islands of Oceania. 

Remarks.—There is reason to suspect that more than one 
species was involved in certain literature references to this species. 
Day (1876, p. 121, pl. 30, fig. 5) gives a high pectoral fin ray (15) and 
lateral line scale count (32 to 34), shows 4 oblique bars on the lower 
lobe as well as the upper lobe of the caudal fin, and 2 rows of red spots 
on the body. These are not characteristic of bensasi. Day listed 
and illustrated 7 spines on the dorsal fin. His account may include 
or represent an undescribed species related to bensasi or to a species 
of the wtiatus group (vittatus, sulphureus, moluccensis). 

The account by Snyder (1907, p. 97, fig. 3) agrees with our speci- 
mens, especially in respect to the number of dorsal spines (7) and the 
absence of bars on the lower lobe of the caudal fin. His description 
of color in life is contradictory to that of Day (1876) and Smith 
(1949, p. 229, pl. 27, fig. 562). Fowler (1933, p. 321, fig. 27) and 
Smith (op. cit.) list the number of dorsal spines as variable, 5 to 8 and 
6 to 8 respectively, but each figure a specimen with 7 spines, the first 
small spine, typical of the vittatus and tragula group, being absent. 

In the Western Indo-Pacific (East Africa, India) this species may 
be represented by another form entirely distinct from that of the 
Philippine-Japan area. U. bensast was not listed by Herre and 
Montalban (1928) nor by Weber and de Beaufort (1931). 

The length of the barbel compared with body length in four size- 
groups is shown in table 6. The data are too meager to conclude 
that any appreciable differences exist. There is no population di- 
vergence indicated from an inspection of the gillraker and vertical 
scale counts separated by geographical localities. 


GOATFISH GENUS UPENEUS—LACHNER 511 


TaBLe 6.—Length of barbel in four size-groups, and the number of gillrakers and 
vertical scale rows, by locality, in Upeneus bensasi 


Barbel length in percent of head length 
Standard length in mm. 


2 AP Sea ee. ee ees (eee || aes 1 7 Fel Se |e eae ID eee eens ee 
LoS ae, ane Ses BE 1 1 Pe ee 2 2 1 2 eee ee ee, 
USL De. 63 SE SEES eee ee jee 2 bly Eee 3 L | _ 2222 |-secee lace See 
GTS SLL Le at a. SEO Ss See eee ee * 2) eee ee ee 2 1 2 Ce ene | ee 1 1 

Gillrakers Vertical scale rows 

Locality 

23 24 25 26 27 29 30 31 
SEIN ESS 3a Sed Eh 2 2 d 2 2 IP eee ee 
<p ES Ret a ees 2 4 ll 4 1 9 3 3 
YS eS aps oe Se SE Se Bp SS Sa eS ene eed ee ee eee 1 2} | seeces= oa eee 


Upeneus asymmeiricus, new species 
Piate 13, Ficure B 
Upeneus tragula Fowler, 1933, p. 339 (in part). 

Holotype-—USNM 154659, a female specimen 76 mm. in standard 
length, collected March 24, 1909, at Pandanon Island, east of Cebu, 
Philippine Islands, by the Albatross Expedition. 

Paratypes—USNM_ 154660, 2 specimens, 73 and 81 mm., taken 
with holotype and having same data; USNM 154661, 1 specimen, 
101 mm., collected April 15, 1908, at Catbalogan, Samar, Philippine 
Islands, by the Albatross Expedition. 

Description.—This description is based on the holotype and three 
paratypes listed above. The counts are recorded for the holotype and 
followed in parentheses by the range of counts for the paratypes. 
Certain characters are compared with other species of the genus in 
tables 1 and 2. 

Dorsal rays VII-i,8 (VII-i,8), the first spine longest; pectoral rays 
ii,11 (ii,11 to ii,12); vertical scale rows 28 (29-30); scale rows above 
lateral line 2 (2-3); scale rows below lateral line 6 (5-6); total number 
of gillrakers 27 (28-29); length of longest raker in longest filament 
1.4 (1.3-1.5). 

Measurements, expressed in thousandths of the standard length, 
are given for the holotype and paratypes in table 7. 

Peritoneum silver brown; preorbital scales present; barbels of aver- 
age length, extend beyond eye but not reaching preopercular margin, 
and barbel length in percent of head length 58 to 66; first dorsal spine 
longest. 

Body slightly slab-sided; jaws nearly horizontal; snout rounded; 
pectoral fins pointed; anal slightly falcate; caudal fin deeply forked. 


512 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


TABLE 7.—Measurements of Upeneus asymmetricus expressed in thousandths of 
the standard length 


Holotype | Paratype | Paratype | Paratype 
Characters USNM USNM USNM USNM 
154659 154660 154660 154661 
ayer ag Jengthy mririt a8 8 coe 8 eee Fe ee 76 81 73 101 
depth (approximate) !-® 2 ts es Es ee EE 3 ene 232 232 
= (approximate)! 2 .-- + 322 228.2 5 £2 145 143 136 144 
ea 

| [i1) 73 1 OE ee Ce eS eee amen a ers Ae 290 287 279 

epth esate ete anes sane hao ee Re 192 189 184 178 
be peduncle 

nage a og Se Ss eh a ee ee ee 279 251 241 228 
TEASE UG DG ise ne en oe re ee eee eee See 96 98 88 94 

Titernrtitienr lenst bony: width=s a ee ee 82 79 69 
Snout, ength as eR ee See eee ee See ee 99 103 104 112 
Orbit, leneth #4 22 Wo fea 222 ane SS 4 78 75 67 
Upper jaw, leneth-. oh ne enna eee 105 101 98 105 
Barbeloteneth-4..- .- 25 ses. Fhe ah he Ry 184 178 195 188 
Spinous dorsal fin, depressed length___-___-_---_------- 214 198 196 203 
Pectoral fin, leng SR EP Re aS ot PR Fe 209 210 200 212 
Pelvic fin, tenet fee Mead ectae petaresetes eee I kas Soe Sele) Aoi tis 210 214 196 197 
Anal fin, depressed length-_---_-..-.-_.-----=----------- 165 178 169 192 
Tip of snout to origin of spinous dorsal fin--_----------- 370 356 368 364 
Tip of snout to origin of anal fin-____--_---_--_---------- 621 615 639 655 


Color in aleohol—Head and body light tan, darker tan above and 
ligher below; a brown, horizontal stripe, its width about one-half 
diameter of eye extends from junction of gill opening with body to 
area just above midbase of caudal fin; a dark brown saddle on caudal 
peduncle just posterior to soft dorsal fin, its width equals least depth 
of peduncle and extends ventrally to lateral line; traces of brown color 
pattern indicating that probably two more saddles existed through 
soft dorsal and spiny dorsal fins; a small, dark spot at origin of spiny 
dorsal and soft dorsal fins and one dorsally on peduncle, at first pro- 
current caudal ray; head and body dorsally above lateral line with a 
“salt and pepper” coloration. 

Spiny dorsal fin with 4 rows of faint tan spots forming horizontal 
bars; soft dorsal with 3 narrow, brown, horizontal bars; pelvic fins 
with traces of 3 or 4 bars; caudal fin with 3 narrow, oblique brown 
bars on upper lobe, the clear interspaces more than twice width of 
bar; lower lobe with 6 or 7 brown oblique bars, more or less spotlike 
and not on the two outermost developed rays (lower lobe on USNM 
154661 malformed and small); areas on caudal fin just above and below 
base with a fine, brownish spot; pectoral and anal fins transparent. 

Named asymmetricus in reference to the unusually reduced number 
of oblique bars on the upper lobe of the caudal fin. 

Geographical distribution—At present, only known by the type 
specimens listed above from the Philippine Islands. 

Remarks.—The number of oblique caudal bars, more than twice 
as many on the lower lobe as on the upper, and their narrow width 
coupled with the absence of the small first dorsal spine, low number 
of pectoral rays, low vertical scale row count, dark lateral stripe, 


-GOATFISH GENUS UPENEUS—LACHNER 513 


dark saddle posterior to soft dorsal, and moderate-to-short barbels 
best characterize this species. 


Upeneus sulphureus Cuvier and Valenciennes 
PuiaTE 13, Figure C 


Upeneus sulphureus Cuvier and Valenciennes, 1829, p. 450 (type locality, Antjer 
Straits of Sundra).—Bleeker, 1877, pl. (8) 393, fig. 4—Smith, 1949, p. 229, 
pl. 28, fig. 563.—Fowler, 1933, p. 330. 

Upeneus bivittatus Cuvier and Valenciennes, 1831, p. 520 (type locality, Coro- 
mandel). 

Mullus subvittatus Temminck and Schlegel, 1843, p. 30 (type locality, Japan).— 
Boeseman, 1947, p. 43. 

Upeneoides sulphureus Bleeker, 1849, p. 63.—Day, 1876, pt. 1, p. 120, pl. 30, fig. 
3.—Herre and Montalban, 1928, p. 103, pl. 3, fig. 1. 

Mulloides pinnivittatus Steindachner, 1870, p. 624 (type locality, Nagasaki). 

Upeneoides belaque Fowler, 1918a, p. 40, fig. 16 (type locality, Philippines). 

Specimens studied —Two hundred seventeen specimens, ranging in 
length from 48 to 132 mm., from the following localities: Zanzibar, 
USNM 12614, 3 specimens, 72 to 79 mm., received from the British 
Museum; East Indies Islands, four USNM collections totalling 10 
specimens; Philippine Islands, 36 USNM collections totalling 200 
specimens; China, USNM 94814, one specimen, 132 mm.; Okinawa, 
USNM 71838, one specimen, 107 mm.; Fiji Islands, Suva, USNM 
66069, one specimen, 91 mm.; Pacific, USNM 83115, one specimen, 
89 mm., Wilkes Expedition. 

Description —Dorsal rays VIII-i,8(48), the first spine minute; pec- 
toral rays 15.8: 15 to 17 (58); vertical scale rows 35.4: 34 to 37 
(44); scale rows above lateral line 3 (10); scale rows below lateral line 
7 (27); total number of gillrakers 29.4: 26 to 32 (43); rakers com- 
paratively long and slender, length of longest raker in longest filament 
averages about 1.1 (6). 

Peritoneum dark, uniform light brown to blackish brown; preorbital 
scales absent; barbels show considerable variation in length and are 
longer in larger specimens (table 8), usually extending beyond pre- 
opercular margin in specimens over 85 mm. in standard length, barbel 
length in these larger specimens in percent of head length 58 to 82 
(73 specimens) ; range of barbel length in percent of head length, all 
specimens, 50 to 82 (84 specimens); third dorsal spine slightly longer 
than second or fourth. 

Color in alcohol—Head and body silvery tan to golden tan, darker 
brown dorsally and light silvery to golden tan below. 

Spinous dorsal with three blackish horizontal bars located near 
base, middle and outer parts of fin, and separated from each other 
by whitish bars; the outer black bar is the most intensely developed 


279126—54——_3 


514 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


and is somewhat blotched; soft dorsal with three dusky to black bars, 
usually less intense than those of first dorsal and separated by three 
light or whitish bars often completely faded; caudal fin with some 
dusky near tips of rays, especially near fork; pectoral, pelvic, and 
anal fins clear. 

Geographical distribution—From East Africa eastward through the 
East Indies, Philippines, and certain island groups of western Oceania 
(Fiji, USNM 66069; New Hebrides, Herre, 1936, p. 210), and north- 
ward to Japan. 


TABLE 8.—Length of barbel in Upeneus sulphureus 


Barbel length in percent of head length 
Standard length in mm. - 
50 | 52 | 54 | 56 | 58 | 60 | 62 | 64 | 66} 68 


70 | 72 | 74 | 76 | 78 | 80 | 82 


— | —— ] ——— |] ——_ |] ——__ ] —____ |] 


41-60... ..Losutiquei.ic i y ft a TE | Li oly, 2060]. SUPOZt S2a saa Ree ae 
Ol=80 oa ee oan ewes ceae re eS | eS Ee ee 2] 3} .4 |---|. 0}. 2) Tee 
SI-100 1s eeree as ert ss | esd are Pe ae VDF (2 1} 2) 4) 3] 2) 3) 1 1 
Over"I00 = 55-35. 25-55255) 2-20) 52-5| cso )aong|=anqjeons]ones A eR Wa | Ral Pp 2 2 


Remarks.—lIllustrations of this species in “‘life colors” show con- 
siderable differences in the development and intensity of the bars on 
the dorsal fins and general body coloration, even those completed by 
recent artists (Smith, 1949, pl. 28, fig. 563; Herre and Montalban, 
1928, p. 103, pl. 3, fig. 1). These differences may be associated with 
state of preservation, size, or perhaps sexual dimorphism. 

The recognition of Upeneus subvittatus Temminck and Schlegel by 
Synder (1907, p. 101) is probably entirely erroneous in view of the 
recent account by Boeseman (1947, p. 43) on the reexamination of 
the type material and on my examination of a specimen collected by 
Jordan and Snyder at Wakanoura, Japan (see ‘‘Remarks” in description 
of tragula). 


Upeneus moluccensis (Bleeker) 
Puate 13, Figure D 


Upeneoides moluccensis Bleeker, 1855, p. 409 (type locality, Amboina).—Seale, 
1914, p. 68, pl. 392, fig. 1—Herre and Montalban, 1928, p. 101, pl. 6, fig. 1. 

Upeneoides dubius Kner, 1865, p. 67. 

Upeneoides fasciolatus Day, 1868a, p. 151 (type locality, Madras). 

Upeneoides sulphureus Day, 1876, p. 120 (in part). 

Upeneus moluccensis Bleeker, 1877, pl. (2) 392, fig. 1—Fowler, 1933, p. 328. 

Upenoides (error) moluccensis Herre and Montalban, 1928, p. 101. 


Specimens studied.—Twenty-one specimens, ranging in length from 
49 to 125 mm. from the following localities: East Indies Islands, 
Borneo, Sandakan Bay, USNM 1388638, 6 specimens, 66 to 74 mm.; 
Java, USNM 72694, one specimen, 82 mm.; Philippine Islands, nine 
USNM collections totalling 10 specimens, taken chiefly by the Alba- 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 


A, Upeneus bensasi (Temminck and Schlegel), USNM 71356, 85 mm., Shimizu, 
Suruga Bay, Japan; B, U. asymmetricus, new species, holotype, USNM 154659, 
76mm., Pandanon Island, Philippines; C, U. sulphureus Cuvier and Valenciennes, 
USNM 145207, 93 mm., Parang, Mindanao Island, Philippines; D, U. moluccensis, 
(Bleeker), USNM 138629, 94 mm., Balayan Bay, Luzon Island, Philippines; E, 
U, vittatus (Forskal), USNM 106850, 147 mm., Iloilo, Panay Island, Philippines. 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 14 


ee qth L ke ee 


= QR 


~~ o® 
Ne el Fr 


A, Upeneus arge Jordan and Evermann, USNM 115685, 213 mm., Canton Island, Phoenix 
Islands; B, U. luzonius Jordan and Seale, USNM 106829, 81 mm., Iloilo, Panay Island, 
Philippines; C, U. tragula Richardson, USNM 145233, 105 mm., Luzon Island, Philippines; 
D, U. oligospilus, new species, paratype, USNM 147995, 124 mm., Tarut Bay, Persian Gulf. 


GOATFISH GENUS UPENEUS—LACHNER 515 


tross Philippine Expedition; Japan, Kagoshima, Satsuma, USNM 
71354, 2 specimens, 106 and 125 mm. (Samoan Islands, USNM 41559, 
2 specimens, 78 and 79 mm., questionable identification). 

_ Description —Dorsal rays VIII-i,8(15), the first spine minute; 
pectoral rays 16.0: 15 to 18 (20); vertical scale rows 34.5: 33 to 36 
(13); scale rows above lateral line 3: (7); scale rows below lateral 
line 7: (8); total number of gillrakers 28.9: 27 to 31 (20); length of 
longest raker in longest filament 1.0 to 1.4 (5). 

Peritoneum uniform light brown to dark brown; preorbital scales 
absent; barbels extend to area between eye and preopercular margin, 
and barbel length in percent of head length 48 to 64 (21 specimens) ; 
second and third dorsal spines about equal in length and these only 
slightly greater than fourth spine. 

Color in alcohol.—Head and body pale to brown above and silvery 
to light tan below; a sharp horizontal, lemon-yellow stripe persists 
on the body in some specimens after about 50 years of preservation 
and extends from eye to area just above midbase of caudal fin. 

The spiny dorsal fin has three dark brown bars separated by three 
transparent to whitish bars, and the tips of the second to fourth 
spines are whitish; soft dorsal with brown bars separated by two trans- 
parent to whitish bars with the tips of the longest rays sometimes in 
white; caudal fin with three to four brownish to dusky oblique bars 
on the upper lobe, the lower lobe clear to dusky, especially near tips 
of rays; pectoral, pelvic, and anal fins clear. 

Geographical distribution.—From India (Day, 1868 and 1876) east- 
ward through the East Indies to the Philippines and possibly Oceania 
(Samoan Islands); from Japan (Kagoshima) southward to Australia 
(Kner, 1865). 

Remarks.—Certain characters given in the account and shown on 
the illustration of Mullus dubius Temminck and Schlegel (1843, p. 
30, pl. 11, fig. 3) can be associated, in part, to those of U. moluccensis. 
The narrow, yellow stripe on the body, the oblique bars only on the 
upper lobe of the caudal fin, and the bars on the dorsal fins as shown 
for dubius are characteristic of moluccensis. Seven spines in the 
spiny dorsal fin were listed for dubius, but a report on the type by 
Boeseman (1947, p. 44) revealed eight. Boeseman found the type 
specimen in a very bad state of preservation and the color and color 
pattern were faded. He recently examined the dentition of the type 
of dubius for me and found the teeth of the jaws to be rather stout 
and large and clearly separated from each other. No teeth were on 
the vomer or palatines. Thus Mullus dubius Temminck and Schlegel 
must definitely be referred to the genus Parupeneus Bleeker (1868, 
p. 344). We may never know if more than one species was involved 
in their description. 


516 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 
Upeneus vittatus (Forskal) 


PiatE 13, Figure E 


Mullus vitiatus Forskal, 1775, p. 31; (type locality, Djedda, Red Sea).—Lacépéde, 
1802, p. 382, pl. 14, fig. 1—Shaw 1803, p. 616, pl. 89. 

Mullus bandi Shaw, 1803, p. 615 (type locality, Vizagapatam). 

Upeneus vittatus Cuvier and Valenciennes, 1829, p. 448.—Bleeker, 1877, pl. (2) 
392, fig. 3—Fowler, 1933, p. 334, fig. 31; 1949, p. 95.—Smith, 1949, p. 228, 
pl. 27, fig. 561. 

Upeneus bitaeniatus Bennett, 1831, p. 59 (type locality, Mauritius). 

Hypeneus vittatus Cantor, 1850, p. 1017. 

Upeneoides vittatus Gunther, 1859, p. 397.—Day, 1876, p. 120, pl. 30, fig. 2.— 
Sauvage, 1891, p. 219 (not pl. 27, fig. 2)—Herre and Montalban, 1928, p. 
105, pl. 4, fig. 1. 

Upeneoides caeruleus Day, 1868b, p. 194 (type locality, Madras); 1876, p. 121. - 

Upeneoides vittatus Klunzinger, 1870, p. 741 (error). 

Upeneoides philippinus Fowler, 1918a, p. 37, fig. 15 (type locality, Philippines). 


Specimens studied.—One hundred fifty specimens, ranging in length 
from 56 to 226 mm., from the following localities: Zanzibar, USNM 
154172, 1 specimen, 67 mm., received from British Museum; Mauri- 
tius, USNM 19956 and 19985, 19 specimens collected by Col. N. Pike; 
East Indies, USNM 72693, 88032, 88033, 145288, and 145290, 6 
specimens from Sumatra, Java, Borneo, and Bouro Islands; Philip- 
pines, 32 USNM collections, 93 specimens collected by the Albatross 
Philippine Expedition, The Philippine Commission, Mearns and 
McGregor; Japan, Okinawa, USNM 71679, 4 specimens, collected 
by the Albatross; Marianas, USNM 124088 and 139854, 7 specimens, 
collected by Frey, McElroy, and Markley; Fiji Island, USNM 
66070, 1 specimen; Samoan Islands, 3 USNM collections, 10 speci- 
mens; Society Islands and Tuamotus Islands (Low Archipelago), 
USNM 89045 and 133844, 3 specimens; Marquesas Islands, USNM 
89750, 6 specimens, collected by the Pinchot Expedition. 

Description.—Dorsal rays VIII-i,8(48) the first spine minute (one 
specimen with 7 spines, the last abnormally small, the first minute and 
typical); pectoral rays 16.1: 15 to 17 (68); vertical scale rows 35.0: 
33 to 37 (49); scale rows above lateral line 3 (13); scale rows below 
lateral line 6.9: 6 or 7 (23); total number of gillrakers 28.2: 26 to 
31 (68); length of longest raker in longest filament averages about 
1.2 (6). 

Peritoneum dark brown to silvery brown (in many of the smaller 
specimens the peritoneum is brownish with a silvery cast); pre- 
orbital scales absent; barbels extend beyond eye but not beyond 
preopercular margin, barbel length in percent of head length 46 to 66 
(74 specimens); third dorsal spine equal to or slightly greater than 
second. 


GOATFISH GENUS UPENEUS—LACHNER 5l-7, 


Color in alcohol_—Head and body light tan to golden tan, darker 
above and lighter tan to silvery below; two to three faint, dark, 
horizontal stripes situated dorsolaterally above middle of body. 

Spinous dorsal fin with two blackish horizontal bars, one on outer 
and one near middle of fin; the outer bar passes through outer third of 
2nd to 5th spines, and is colored an intense black; a whitish bar be- 
tween these bars; soft dorsal with three dusky to black bars or marks, 
a short mark near the posterior basal area, a long horizontal bar at 
middle of fin and a narrow dusky tip on the Ist to 3rd rays; caudal 
fin with oblique, dusky to black bars, 3 or 4 on the upper lobe and 2 or 
3 on the lower; sometimes the tip of the longest (outer) rays of the 
upper lobe are slightly touched in black (these I have considered as a 
bar in my counts); outer bar on lower lobe with more intense black 
and twice as wide as other bars on caudal; this particular bar never 
at tip of lower lobe (only one specimen with tip of the lower lobe 
touched in blackish, see table 9); two nearly horizontal bars extend 
from near fork of caudal to areas just above and below midbase of 
fin and were not included in counts of the total number of oblique 
bars of the caudal fin; pectorals, pelvics, and anal fins transparent. 


TaBLe 9.—Relationship of number of bars on caudal fin with increase in body 
length, and the length of chin barbel by four size-groups in Upeneus vittatus 


Number of bars on lobes of 
caudal fin 


Total length in mm. 


Upper 3 | Upper 4 | Upper 4 
Lower 2 | Lower 2 | Lower 3 


fen en ee a oe oe a oa ae one ee eee eo rena ne Po Nee ee een ee 
ae ok hn nn naa nnn d Jes see aye es ss oe eee nae asad 2 5 2. eae cdece cs 
ASN ee Se ee ee ae eee ae eee 2 5 1 
LE Lae oe a ppl a a le A — tie tS 5 6 ee 
Dip AGO ete ee ooo lat ae eee oe BIS. Ee 5 Oe eee ee ees 
PUNE a as Soa a ee sa a eo SE EE ees “SEF WS 


PRA a ee ee. _ eee OPA Ae VRS PARR? SP Ne To R82 ohh ot YS 2 SOT. Bees | 
CSlLe 2 ae eee eee eee 2 (ee 2 | ee pm ie eee 3 1 2 4 3 1 Oh ee 
CUS ae ee See ao ead | a. Re ee 1 i 5 4 a 3 Oo ascot asense 
Oo ee ee ee eee ee 4 | ee 3 6 6 3 8 | See 2 1 


Geographical distribution.—A widely ranging species, occurring from 
the Red Sea eastward through the East Indies and islands of Oceania 
to the Low Archipelago, and reported from Japan southward to 
Australia. Fowler (1933, pp. 334-335) has given an almost complete 
list of references, but many of these include only lists of species and 
such records cannot be accurately appraised. This species has not 
been reported from the Hawaiian and Johnston Islands. 


518 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Remarks.—There is no evidence of any change in barbel length with 
increase in body length from the statistics presented in table 9. The 
relationship of the number of bars on the caudal fin with increase in 
body length is also shown in table 9. At most, one more bar is some- 
times formed on the upper lobe in the larger specimens and this bar 
is merely a touch of blackish at the tips of the outer rays. Only one 
specimen had such a mark on the lower lobe. 


TABLE 10.—Certain counts of Upeneus vittatus, by locality 


Gillrakers Lateral line scales 
Locality 
26 apf 28 29 30 31 33 34 35 36 37 
EN TAD NYS oe oe nome eee di eS Nap fa | eae | of ps pM ae pee ee i | 
Mauritius 2¢)- 1 Gaye. eats Seay 3 45\3. 23 Volt $e. jee Siege. 3 3 2 tae 
ast indies. 36 Jae Pee oa ee | See C1 SR [ae eee eee 1 digloacan pease 
Philippinesiat) ee ee eee 1 Walls Whe. eles t| RAs 9| u Co ee ee 
ANAND. oe see eee ao soa See ae 1 2 1 |=-=-.4|---=£.|--=.-.|. " Ue ee ee 
Marianas 4624 2 O39) 6 obese. ee 3 3 lupe 2 dl eos | ee 1 1 Uy ee 
Te Ae An Sieh RR a | a|| og | PE See he a 1 [2a See 
Samoan’! steve. pes eee ee el coe cue _ ewes 5 (a See eel eee e 1 2 5ileese2- 
Society,,.CusmotuS: 222-222-2022 | eee | ba eo 1 | era a Deane eo | ae 
Marquesas 14: uarheciri et sai At 22 2 sh eee | be ee ne ee -Pee eee Pees ee 3 2 1 


Very little intraspecific differentiation was found in the subfaunal 
areas from an analysis of color and color pattern, body proportions, 
and meristic counts. In table 10 the statistics of the number of gill- 
takers and vertical scale rows are segregated by locality, from the 
east African area eastward. Some of these localities represent sub- 
faunal areas of the Indo-Pacific region, and although a maximum 
separation of about 12,000 miles exists between the extreme localities, 
the data does not suggest any significant population divergence. 


Upeneus arge Jordan and Hvermann 


PuatE 14, Fiaure A 


Upeneus arge Jordan and Evermann, 1903, p. 187 (type locality, Honolulu); 
1905, p. 264, pl. 39.—Fowler, 1928, p. 227, pl. 19,c; 1931, p. 336; 1938, pp. 
224, 285; 1940, p. 777; 1949, p. 96.—Jenkins, 1902, p. 456.—Schultz, 1943, 
p. 128.—Snyder, 1904, p. 527. 

Upeneoides arge Fowler, 1922, p. 83.—Jordan, Evermann, and Tanaka, 1927, 
p. 674.—Jordan and Jordan, 1922, p. 52. 

Specimens studied.—Hleven specimens, ranging in length from 164 
to 250 mm.: Hawaiian Islands, USNM 50667 (holotype), 17999, 
52817, 55100, 83358, 83449, 88194, and 151524, 9 specimens; Phoenix 
Islands, USNM 115685, 2 specimens. 

Description.—Dorsal rays VIII-i,8(11), the first spine minute, 
pectoral rays 13.9: 13 to 14 (12); vertical scale rows 37.3:37 to 38 
(10); scale rows above lateral line 3 (6); scale rows below lateral line 


GOATFISH GENUS UPENEUS—LACHNER 519 


7 (6); total number of gillrakers 22.3:21 to 24(11); length of longest 
raker in longest filament 1.5 to 1.7 (4). 

Peritoneum transparent; preorbital scales absent; barbels extend 
beyond eye to about the preopercular margin and barbel length in 
percent of head length 64 to 74 (10 specimens); second dorsal spine 
equal to or slightly longer than third. 

Oolor in alcohol.—Head, body, and barbels light tan to pale above, 
lighter on chin and belly; two very faint horizontal stripes on body, 
dusky in color, and about as wide as pupil; one stripe is located dorso- 
laterally and the other medially. 

Spinous dorsal with some dusky on outer membrane, remainder 
clear or faintly dusky, soft dorsal with three narrow, transverse bars 
colored diffuse brown; pectoral, pelvic, and anal fins clear; caudal fin 
with 6 oblique brown to black bars on upper lobe and 5 on lower 
lobe; lobes tipped in brownish black; the outer two bars on lower 
lobe about twice as wide as those on upper lobe; two nearly horizontal 
streaks near fork of caudal, just above and below median line. 

Geographical distribution.—The Hawaiian Islands and various island 
eroups of Oceania. Our specimens are from the Hawaiian and Phoenix 
Islands. Fowler (1928, p. 227) reports the species from the Palmyra, 
Caroline, and Gilbert Islands. 

Remarks.—Fowler (1928, p. 227) conjectured that U. arge “may 
eventually be found inseparable” with U. vittatus. He pointed out 
that vittatus has slightly smaller scales than arge but he overlooked 
such trenchant characters as the differences in the color of the peri- 
toneum, number of pectoral fin rays, and number of gillrakers. 


Upeneus luzonius Jordan and Seale 
PLate 14, Figure B 


Upeneus luzonius Jordan and Seale, 1907, p. 25, fig. 9 (type locality, Cavite).— 
Weber and de Beaufort, 1931, p. 372.—Fowler, 1933, p. 325, fig. 28. 

Upeneus sundaicus Evermann and Seale, 1907, p. 88. 

Upeneoides luzonius Seale, 1910, p. 279.—Herre and Montalban, 1928, p. 97, pl. 
1, fig. 1. 

Specimens studied.—Sixteen specimens from the Philippines ranging 
in length from 25 to 119 mm., from the following Islands: Panay, 
USNM 106829, 102649, 106793, 106846, and 154201, 7 specimens; 
Luzon, USNM 53067 (cotypes) and 138658, 5 specimens; Linapacan, 
USNM 138659, 3 specimens, USNM 154200, 1 specimen. 

Description.—Dorsal rays VIII-i,8(11), the first spine minute; 
pectoral rays 14.1:14 to 15 (10); vertical scale rows 31.3:31 to 32 
(8); scale rows above lateral line 3 (5); scale rows below lateral line 
6 (5); total number of gillrakers 20.4:19 to 22 (9); length of longest 
raker in longest filament averages about 1.5 (5). 


520 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Peritoneum transparent; preorbital scales present; barbels long, 
extend to preopercular margin, barbel length in percent of head length 
62 to 72 (10 specimens) ; second dorsal spine longest. 

Color in alcohol_—Head and body tan to dark brown on upper half 
of body, lighter below; smaller specimens lighter tan than larger 
specimens; a dark brown stripe extends from snout through eye to 
area just above midbase of caudal fin, being below the lateral line 
on anterior part of body and above lateral line on posterior portion; 
width of lateral stripe about three-fourths diameter of eye; lateral 
stripe most pronounced in adults and more conspicuous than illus- 
trated by Fowler (1933) or Herre and Montalban (1928); body with 
three dark brown saddles, their width about 1 to 2 times greater than 
diameter of eye; first saddle passes through midbase of spinous dorsal 
fin, the second through midbase of soft dorsal, and the third passes 
over caudal peduncle just posterior to soft dorsal fin; saddles extend 
ventrally to the lateral line and are more conspicuous in smaller 
specimens; the two anteriormost saddles almost completely faded in 
the larger specimens. 

Spinous dorsal dusky on upper half, three very faint, transverse, 
dusky bars in specimens about 50 mm. in length; soft dorsal with 2 
faint, dusky, transverse bars in smaller specimens and 3 in the larger 
ones; pectoral fin with about 3 to 5 barely discernible vertical bars on 
one adult specimen, fins of other specimens completely clear; pelvic 
fin with a faint brownish blotch or bar in specimens about 50 mm. in 
length, fins of largest specimens clear; anal fin with a faint dusky bar 
in smaller specimens, completely clear in larger specimens; caudal 
fin with 2 to 7 oblique, dusky-to-brown bars on each lobe, the bars 
increasing in number with increase in length (table 3); the bars on the 
lower lobe broader, wider than the clear interspaces, and more in- 
tensely developed than those of the upper lobe; bars on upper lobe 
almost as wide as clear interspaces; bars on fins in varying degrees 
of intensity, the caudal bars being most evident. 

Geographical distribution.—East Indies (Seale, 1910; Herre and 
Montalban, 1928) and Philippine Islands. 

Remarks.—The differences in the color pattern between Fowler’s 
illustration (1933, fig. 28) and that of Herre and Montalban (1928, 
pl. 1, fig. 1) are undoubtedly associated with the state of preservation 
and the sizes of specimens used for the illustrations. Fowler indi- 
cated his specimen was a ‘‘young’”’ one. The low number of caudal 
bars, fairly conspicuous bars on the dorsal fins, and well-developed 
saddles, as he has illustrated, are characteristic of the smaller sizes. 
Herre and Montalban illustrate about 7 bars on each lobe of the caudal, 
which is characteristic of specimens about 100 mm. in length (table 3). 


521 


GOATFISH GENUS UPENEUS—LACHNER 


This species is closely related to U. tragula, and the smaller speci- 
mens of each species are easily confused with each other. The geo- 
graphic range of tragula completely overlaps that of luzonius and 
both have been collected together. The young of tragula often have 
a moderately developed saddle on the caudal peduncle, generally 
lack the spotting characteristic of the adults, and the color pattern 
of the fins may be nearly obscure; consequently, these specimens 
superficially resemble luzonius. The characters listed in table 11 are 


most helpful in distinguishing between these species. 


TasBiLe 11.—Characters distinguishing Upeneus luzonius and U. tragula 


Character 


Pectoral fin rays. 


Total number of gill- 
rakers. 

Barbel length in per- 
cent of head length. 

Brown spots on head 
and body. 


Dark brown saddles on 
body.? 


Pigmentation of: 
(a) dorsal fins. 


(b) pelvics and 
anal fins. 


(c) caudal fin. 


luzonius 


92 percent with 13 or 
fewer, range 12 to 14. 
20.4 (19 to 22).! 


66.6 (62 to 72). 


Always absent. 


3 in young, 2 or 1 (the 
posteriormost one) in 
adults; sometimes all 
ar: almost completely 
faded. 


2-3 faint dusky bars on 
each but often com- 
pletely faded. 


Clear in adults; each with 
a faint transverse bar 
in small specimens. 

Dark brown oblique bars 
nearly uniformly nar- 
row and elongate; de- 
veloped with almost 
equal intensity on each 
lobe; usually occur- 
ring in equal numbers 
on the lobes; 1 bar 
more per lobe at a 
given length (table 3). 


1 The mean is followed by the range of variation in parentheses. 
2? All descriptions of color and color pattern refer to preserved specimens. 


tragula 


All with 14 or 15. 
22.8 (21 to 25). 
59.2 (52 to 68). 


Usually well developed and 
numerous, sometimes faint 
to obscure. 

Faint to moderately devel- 
oped saddle over caudal 
peduncle in young and 
juvenile specimens, often 
inconspicuous or entirely 
faded in adults. 


Each almost always tipped 
in a large blackish irregu- 
lar blotch, remainder of 
fin blotched with black, 
white, or clear. 

Almost always with dark 
brown spots arranged in 2 
to 3 rows. 

Dark brown, oblique bars, 
elongate anteriorly becom- 
ing oval-shaped posteri- 
orly, especially outer two 
bars on lower lobe; bars on 
lower lobe conspicuously 
more intensely developed; 
usually 1 more bar on the 
lower lobe; 1 bar less per 
lobe at a given length 
(table 3). 


522 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Upeneus tragula Richardson 
PuiaTe 14, Figure C 


Upeneus tragula Richardson, 1846, p. 220 (type locality, Canton).—Fowler, 1933, 
p. 339 (in part). 

Upeneoides variegatus Bleeker, 1849, p. 64 (type locality, Batavia). 

Upeneoides kiushiuana Déderlein, in Steindachner and Déderlein, 1884, p. 22 
(name only; type locality, Kagoshima). 

Upeneus subvittatus Snyder, 1907, p. 101. 

Upeneoides tragulus Snyder, 1912, p. 503. 

Specimens studied.—One hundred eighty-eight specimens, ranging 
in length from 29 to 227 mm., from the following localities: Zanzibar, 
USNM 12614, 1 specimen, 83 mm., received from the British Museum; 
East Indies Islands, USNM 72695, 145263, and 145624, 6 specimens, 
79 to 188 mm.; Philippine Islands, 64 USNM coilections, 115 speci- 
mens, most taken by the Albatross Philippine Expedition; China, 
USNM 9128, 145265, and 148413, 3 specimens; Japan and Okinawa, 
9 USNM collections, 13 specimens; Palau Islands, USNM 154202, 
47 specimens collected by Eugenie Clark, July 1949; Australia (New 
South Wales), USNM 59957 and 82984, 3 specimens. 

Description.—Dorsal rays VIII-i,8(35), the first spine minute; 
pectoral rays 13.0: 12 to 14 (77); vertical scale rows 30.0: 28 to 32 
(48); scale rows above lateral line 3 (9); scale rows below lateral line 
6 (37); total number of gillrakers 22.8: 21 to 25 (57); length of 
longest raker in longest filament averages about 1.5 (5). 

Peritoneum transparent to silvery; preorbital scales present; barbels 
extend to area between eye and preopercular margin, barbel length 
im percent of head length, 52 to 68 (58 specimens); third dorsal spine 
slightly longer or about equal to fourth. 

Color an alcohol.—Dorsal portion of head and body pale or dusky 
to tan; chin and belly lighter; barbels pale; a tan to blackish brown 
horizontal stripe extends from tip of snout, through eye, along middle 
of body to base of caudal fin; width of stripe somewhat variable, 
about 0.5 to 1.0 in diameter of eye; chin, cheeks, operculum, and 
body below lateral line with numerous circular to irregularly shaped 
spots, colored tan to dark brown or dusky; these spots are more 
iregular, larger, blotchlike and less intense on some of our larger 
specimens over 160 mm.; spots sometimes faint or completely absent. 
(Specimens of the large collection recently taken in the Palau Islands 
lack well-developed spots almost entirely, whereas many specimens 
taken nearly a half century ago by the Albatross Expedition in the 
Philippines have well-defined spots. These island groups are contig- 
uous, geographically, and such color variations may be associated 
with different methods employed in collecting and preserving rather 
than racial differences.) Dorsal portion of head and body usually 


GOATFISH GENUS UPENEUS—LACHNER 523 


finely speckled with brown; lower sides of body with 8 to 10 faint, 
dusky blotches or bars, variable in size and often completely obscure; 
an inconspicuous dusky to brown saddle, sometimes obsolete, just 
posterior to soft dorsal fin, its width about two-thirds length of base 
of soft dorsal. 

Spinous dorsal fin with large, brown-to-blackish, irregular spot on 
outer third, a clear or whitish spot anteriorly on basal third followed 
by a large, brownish-black, irregular spot midbasally, remainder of 
fin transparent; soft dorsal transparent with three brown to black 
marks, an outer spot, a horizontal stripe near middle, and a short 
stripe near base on anterior third; pectoral fin transparent with one 
circular, brown to dusky spot near base; pelvic fin transparent with 
6 to 9 circular, brown to black spots arranged in 2 to 3 rows, often 
completely faded; anal fin transparent with 2 to 3 bars (or spots, 
depending on condition of fading), brown to blackish colored and 
almost parallel to base; caudal fin transparent with 2 to 6 oblique 
bars on the upper lobe and 2 to 7 on the lower lobe, colored brown 
to black, the number of bars increase with increase in length (table 
3); lower lobe usually has one more bar than the upper lobe, and 
coloration of bars more intense; the caudal bars first appear as spots 
which become oval shaped (specimens under 50 mm.) and then 
elongate to form the definitive oblique bars. 

Geographical distribution—Represented in the U. S. National 
Museum collections from East Africa, eastward, in the East Indies and 
Philippines to the Palau Islands and from southern Japan to New 
South Wales, Australia. Although abundant in the Philippine 
Islands (note specimens studied above; Herre and Montalban, 1928, 
p. 99; Fowler, 1933, p. 339; and Weber and de Beaufort, 1931, p. 368), 
this species apparently becomes rare eastward, in certain islands of 
Oceania, and has not been reported for most of the island groups. 
In addition to the excellent collection made by Eugenie Clark from the 
Palau Islands, USNM 154202, Herre (1935, p. 165) also listed it. 
Seale (1935, p. 362) listed specimens from the Solomon and Samoan 
Islands, all of which were small. Herre (1936, p. 209) also reported 
on two small specimens from the Solomons, but his description was 
based on Philippine material. Jordan and Seale (1906, p. 273) 
reported it, too, from the Samoan Islands. Schultz (1943) did not 
collect it in either the Phoenix or Samoan Islands, nor was it taken by 
the recent intensive collecting by Schultz and others in collaboration 
with the U. S. Navy project in the Marshall Islands (1946-1947). 
Superficial examinations, particularly of the small specimens, could 
easily lead to misidentifications, especially with such forms as U. 
luzonius and vittatus. Specimens reported from the Solomons and 


524 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Samoan Islands should be reexamined. U. tragula has not been 
reported from the Hawaiian faunal area. 

Remarks.—The change and variation in length of barbel with in- 
crease in body length was investigated (table 12) but no appreciable 
difference was found. 

This species appears constant in body proportions, meristic counts, 
and coloration over its range. Counts of the number of gillrakers and 
vertical scale rows were segregated by locality (table 12) and these 
data do not even suggest any population divergence. 

U. tragula is most closely related to oligospilus and luzonius and 
compared with them in their descriptive accounts. Table 3 shows the 
relationship of the number of oblique bars on the caudal fin with 
increase in body size in these three species. 


TaBLE 12.—Number of gillrakers and vertical scale rows, by locality, and length of 
chin barbel in four size-groups in Upeneus tragula 


Number of gillrakers Number of vertical scale rows 
Locality 
21 22 23 24 25 28 29 30 31 32 
ZANT A Se 8 8 bd a ae ed ee I ee eee ee eee ee ee | 2 een eee ee 
Hast Indies: 2-4. -.202Le. Ane toa | Me EO SEES pas) 1; 9 mee et Eee el ee 2 2 1 1 
Philippines 222-2 sao 2 Sa 3 4 21 Pn eet 2 7 9 6 2 
inal. 2525) sss eC eo aah eee Pts oes Lee it Za ss a eee PLES 
AE 2 ee ee ee ee ee ee 3 @ | i Sa Gee 1 2 i. 
Palawiis ti Bor ts 2 EU Rae) ADIT EL Sea 6 3 2 Se 3 3 2 1 


Standard length in mm. 


41-609 Se EEA ed ered re ND oe Se Fel ee SE AT Dees. ce FP 3 22-2. be Bees 1 
Lip ES ee SMa SP Se patie. we i nae ate slip aes i eect 1 oe | ie |p ee fa eet 2 Bh et 
SEAN OO. M22 shee ors Sart Sa ce ge So Be ele wees Ds see. 6 3 3 1 ee SS 
Greadrer Chan 000 ace ee ee ee ee ee 3 3 3 8 5 2 | secaee | eee 


A specimen of U. subvittatus Snyder (1907, p. 101), Stanford 
University Natural History Museum No. 20156, collected by Jordan 
and Snyder at Wakanoura, Japan, is without question U. tragula. 
The following counts for this specimen almost all fall on the modes of 
the frequency distributions given above for tragula: pectoral fin 
rays 28, gillrakers 2, 4+1-++11, 5 totaling 23, barbel length in head 
length 58 percent, vertical scale rows 29. Other important characters 
as the silvery peritoneum, dark lateral stripe on body, dark dorsal 
saddles, and oblique bars on caudal fin are also identical to those of 
tragula. Snyder gave a “length” of 175 mm. in contrast to my 
measure of 228 mm. (standard length) for the above specimen. 
This discrepancy is not explainable. 


GOATFISH GENUS UPENEUS—LACHNER 525 


Upeneus oligospilus, new species 
PratE 14, Figure D 
Upeneus tragula Blegvad and L¢gppenthin, 1944, p. 135, pl. 7, fig. 3. 


Holotype—USNM 153988, a female specimen 115 mm. in standard 
length, collected April to June, 1948, at Tarut Bay, Ras Tannura, 
Persian Gulf, by Donald S. Erdman. 

Paratypes—USNM 147995, 11 specimens, 70 to 160 mm., taken 
with the holotype and having the same data. 

Description.—This description is based on the holotype and para- 
types listed above. The counts are given for the holotype, followed 
in parentheses by the average and range of counts taken from the 11 
paratypes. When counts for the paratypes are identical with those 
of the holotype, only one number is given. Certain characters are 
compared with other species of the genus in tables 1-3. 

Dorsal rays VIII-,8 (VIII-,7.8: VIII-i,7 to i,8), the first spine 
minute; pectoral rays 14 (13.6: 13 to 14); vertical scale rows 30 (30.2: 
29-31); scale rows above lateral line 3; scale rows below lateral line 6; 
total number of gillrakers 23 (22: 20-23), rakers short and blunt, 
length of longest raker in longest filament 1.5 to 2.3. 

Measurements, expressed in thousandths of the standard length, 
are given for the holotype and paratypes in table 14. 

Peritoneum transparent, slightly dusky in certain areas of some 
specimens; preorbital scales present; barbels of average length, extend 
beyond eye but not to margin of preopercle, barbel length in percent 
of head length 50 to 64; fourth dorsal spine equal to or slightly smaller 
than third. 

Margins of pectoral fins round, spiny dorsal fin round, soft dorsal 
and anal slightly falcate, caudal fin deeply forked. 

Color in alcohol—Head and body pale with dusky to blackish 
blotches of irregular shape and size; body darker above, with scattered 
dusky pigmentation, and lighter below; a dark stripe from tip of 
snout through eye to area just above midbase of caudal fin; diffuse 
dusky spots on snout and cheeks and a few on sides of body in some 
specimens but absent on belly; light or silvery spots on scales of body, 
the diameter about one-third to one-half vertical length of scale; a 
weakly developed, dark saddle just posterior to soft dorsal fin, nearly 
obsolete in most of the specimens. 

Outer third of spinous dorsal with dense black blotch, remainder of 
fin blotched in black, whitish or clear; soft dorsal with an irregular 
black spot near tip, a black spot or incomplete bar near base and some 
scattered, dusky spots; pectoral fin with a small, black spot near base, 
remainder transparent; pelvic fin with 2 to 3 rows of circular, black 
spots, about 6 in all; anal fin with 2 faint, elongate dusky marks, 


526 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 
remainder of fin clear; caudal fin with black, oblique bars on both 
lobes, varying from 2 on the upper and 3 on the lower in the smallest 
specimen and 4 bars on both lobes in the larger specimens (table 3); 
bars on the lower lobe more distinct and slightly wider than those of 
the upper; a diffuse, blackish, irregular, and broken spot near midbase 
of caudal fin. 

Named oligospilus in reference to the faint, scattered, dusky spots 


on the sides of the body. 


Geographical distribution. — Known from the Persian Gulf and proba- 
bly the Gulf of Oman (Blegvad and L¢ppenthin, 1944, p. 136). It 
may occur more widely, especially along the Indian coast, than is now 
known. 

Remarks.—This species is related to luzonius and tragula; con- 
siderably more to the latter species. It differs from tragula chiefly in 
the reduced number of oblique bars on the caudal fin (table 3) and in 
the reduced number of dark spots on the sides of the body, which are 
decidedly more diffuse and larger, and absent on the belly. It differs 
from tragula also in certain other characters, some of which are difficult 
to measure for statistical inspection. Of these it is worthy to mention 
the deeper, more robust body, shorter fins, especially the pectorals 
and pelvics (see table 13), slightly longer head, and higher modal 
count of the pectoral fin rays (table 1). 


TaBLE 13.—Coloration in life of Upeneus oligospilus and of U. tragula 


Item 


Head and body. 


Horizontal stripe. 
Spots on head and body. 


Spiny dorsal fins. 


Caudal fin. 


1 Blegvad and L¢ppenthin (1944) 
2 Herre and Montalban (1928) 


U. oligospilus! 


(Persian Gulf) 

Head brownish dorsally; 
side of body below lat- 
eral stripe with bluish 
green cast; abdomen 
silvery with red tinge. 

Orange. 

Seattered, irregular, 
vivid, vermilion spots, 
quickly fading in al- 
cohol. 

Vermilion bars. 


Vermilion 
rower. 


bars; nar- 


U. tragula? 


(Philippines) 
Grayish yellow; body ven- 
trally white flushed with 
roseate. 


Dusky to brown. 

Conspicuous uniform 
brown spots; sparingly 
to thickly spotted and 
persist in preservation. 

Upper third black with 
circular, yellow spots, 
and lower portion with 
two dusky bars. 

Dusky bars; wider. 


GOATFISH GENUS UPENEUS—LACHNER 527 


Blegvad and Lgppenthin (1944, p. 135) attributed the differences 
in color of their specimens, when compared with the descriptions of 
Weber and de Beaufort (1931, p. 368) and Day (1876, p. 121), as 
apparently due to the preserved state of the material. This may have 
been so in part, but it is now realized that two species were involved. 
A description and figure of the color in life of oligospilus is given by 
Blegvad and Lgppenthin (op. cit., pl. 7, fig. 3) in their account of U. 
tragula, and when it is compared with the color in life of tragula given 
by Herre and Montalban (1928, p. 100, pl. 2, fig. 2) notable differences 
are found. There are some obvious errors in both illustrations, such 
as the diagrammatic arrangement of the bars on the fins of tragula by 
Herre and Montalban, as well as the excessively uniform distribution 
of the spots on the body and the omission of the dark brown saddle 
just posterior to the soft dorsal fin. The illustration of Blegvad and 
Lgppenthin appears to have much of an artist’s touch but lacks details. 
No pigmentation is shown on the outer portion of the spiny dorsal fin 
by them, but in preserved specimens this area has a most conspicuous 
black blotch. The salient differences of color in life between oligospilus 
and tragula as recorded by the above authors are summarized in 
table 13. 


TasB_e 14.— Measurements of Upeneus oligospilus expressed in 
thousandths of standard length 


Characters Holotype} 11 Paratypes! 

Sisndard length amin 2. — ge oes - 2228 $- Se ae sess. = tap lees 115} 105 (70-160) 
Body: 

GOCE Sa Ge5- Hae Se te Se eee Sonn eee by ey a ee eee See ane ee ee eee 238 241 (213-261) 

SUVaT tl kh ie ee eee ee ee ee ee ee ee ee ee eee 157 | 157 (136-178) 
Head: 

I ESES 2 [oa PO le a nce e y  we ee Sn e OR S rae 320 | 306 (280-323) 

CIEE ie et eo ad os ee ae PE See ee RE ee ed nt 195 203 (184-241) 
Caudal peduncle: 

EO Meee ete eee ee ane ee See eee ee ee | eh OE 230 260 (247-277) 

RESTSTRALE DE ete es aa es Rg de Se ee ed tS eee See ee 102 101 (93-107) 
Tnpererpiia: least bony width <<<.) . 2s. 225245 seers se es 82 78 (74 81) 
Silent, Liste Milos Ge Sak eee Red allan a diete fined nek odleek aed leaeaete 130 | 117 (113-131) 
Orhiteene thal OF. . moe we he RK ae eee Eh cee bee) 71 69 (59 78) 
Mi Dpia aie eUtitloms Mere ncone seo se eae ee ne ao eee eee eee 137 129 (119-139) 
Bebe leno nek sso 26 Fe he ES 8 eo Be ee eh nd eel epee bbe 207 179 (152-204) 
Spinous aorsalin, depressed leneth. ..- 9-22 t5_ 22 oot ec ne ee 210 | 214 (194-236) 
Pecranauiine Iencpny a. bo seo- chee Ree US ba eee te ton eS eds 195 202 (167-217) 
eRe PTT eGLO RUE sco Soe ae Re lens oe ees ie Set ee nae 197 197 (160-216) 
Ampletin~ depressed den pth 205 _ . syssas S82 eee eee ces - ee bbe ea aces 174 176 (166-196) 
Pip ersnout tO Origin Of spinous dorsal fins -_*-22- 2222259. 22 (2 oo rar eso oles 378 | 375 (366-390) 
‘imot snout to origin of anal fin. -...-.-2-2.--.s es Je es ed. 662 | 648 (610-697) 


1In paratypes the average values are followed by the range of variation in parentheses. 


TaBLe 15.—Length of pelvic fins in two species of Upeneus, expressed as 
a percentage of the head length 


Percent 
Species 
50 | 52 | 54 so | 38 60 | 62 | 64 | 66 | 68 | 70 | 72 | 74] 76} 78 | 80 | 82 
Obrospilus. _-+ =< 2-229. nee lS 4 eee ee 5 Uf ar A aT Ra | TU VPM | a Pe [ee 


528 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Upeneus parvus Poey 


Upeneus parvus Poey, 1853, p. 226 (type locality, Cuba).—Poey, 1852, pl. 17, 
fig. 4—-Norman, 1922, p. 534.—Longley and Hildebrand, 1941, p. 142. 
Upeneoides parvus Stahl, 1882, pp. 76, 162. 

Specimens studied.—Six specimens from the Western Atlantic 
ranging in length from 68 to 119 mm., from the following localities: 
Cuba, USNM 37576, collected by F. Poey, 1885, 2 specimens; Puerto 
Rico, Anasco Bay, USNM 128263 and 144555, collected by V. Barnes, 
Jr., 1943-1944, 3 specimens; Tortugas, Fla., USNM 92051, collected 
by W. H. Longley, 1931, 1 specimen. 

Deseription.—Dorsal rays VII-i,8, the first spine longest; pectoral 
rays 15.5: 15 to 16 (6); vertical scale rows 37.2: 36 to 38 (4); scale 
rows above lateral line 3: (4); scale rows below lateral line 6.6: 6 to 7 
(3); total number of gillrakers 27.4: 26 to 29 (6); length of longest 
raker on longest filament about 1.2 (4). 

Peritoneum light, transparent to slightly silvery; preorbital scales 
present; barbels extend to posterior margin of preopercle; barbel 
length in percent of head length 62 to 78 (6 specimens). 

Color in alcohol.—Head and body light tan. One specimen shows 
evidence of a light colored median stripe on body and possibly a finer 
one below it. Caudal fin with dark, oblique bars, more pronounced 
on lower lobe; 3 bars on the upper and lower lobes in the smaller speci- 
mens and 4 or 5 on each lobe of the larger ones. Traces of bars or 
marks on the spinous and soft dorsal fins. Remainder of fins clear. 
No photograph was included because of the poor condition of the 
specimens. 

Geographical distribution.—Known from the following localities in 
the Western Atlantic, where it is apparently rare: Cuba, Tobago (Nor- 
man, 1922), Puerto Rico, and Tortugas. 

Remarks.—In Fowler’s description of Upeneus phillipsi (1918b, p. 
5, fig. 1, type locality, Corson’s Inlet, Cape May County, New Jersey) 
he suggested that it may be “allied, if not identical” with Upeneus 
parvus. [ixamination of the holotype of U. phillips: Fowler revealed 
this species to be a juvenile specimen, 52 mm. in standard length, 
of Mullus auratus Jordan and Gilbert. 

Critical generic and specific characters were inaccurately recorded 
by Fowler. He lists and figures 8 spines in the first dorsal fin; states 
that the dentition of the upper jaw consists of at least a row of low, 
simple teeth, and that the vomer and palatines have fine teeth; lists 
30 scales in the lateral line to base of caudal fin and 16 pectoral rays. 
I find 7 spines in the first dorsal fin, no teeth on the upper jaw and those 
of the vomer and palatines coalesced to form a palatal tooth patch 
which is characteristic of the genus Mullus, 34 or 35 vertical scale 
rows along the lateral line to hypural base, and 15 pectoral rays for 
each fin. 


GOATFISH GENUS UPENEUS—LACHNER 529 


The following characters were also recorded from the type specimen: 
total number of gillrakers, 22; barbels extend slightly beyond margin 
of preopercle; peritoneum light colored to slightly dusky; 2 barlike 
marks on spinous dorsal fin and 3 on soft dorsal; caudal fin with 3 dark 
oblique bars on each lobe (tips of lobe now broken off) ; a salmon-col- 
ored stripe on body along lateral line and prolonged through eye with 
coarse flecks of salmon color on the lower portion of sides of body 
(observed in life by collector, R. J. Phillips). 

The characters given above clearly place U. phillips: in the 
synonymy of Mullus auratus. 


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LachPEDE, BERNHARD G. DE La 
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LoneLey, Witu1aM H., and HinpEBRAND, SAMUEL F. 

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532 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


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U. S. GOVERNMENT PRINTING OFFICE: 1954 


PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


Tssued 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1954 No. 3331 


STUDIES IN NEOTROPICAL MALLOPHAGA, XI: 
BIRD LICE OF THE SUBORDER AMBLYCERA, GENUS 
DENNYUS NEUMANN 


By M. A. Carriker, Jr. 


Dennyus Neumann, a genus of Amblycera found only on the swifts 
(Apodidae), is a relatively small group, 13 valid species being listed by 
Hopkins and Clay in ‘“‘A Checklist of the Genera and Species of 
Mallophaga,” 1952. Very little concerning the genus has appeared 
in literature. Apparently few specimens have been collected, owing, 
I suspect, to the difficulty of securing their hosts. 

Of the 13 recognized species, 3 are from North American hosts, 2 
from Central America, 1 from South America, 1 from Jamaica, 1 from 
Europe, and 5 from Asia and the East Indies. 

Harrison (Genera and Species of Mallophaga, 1916) listed six valid 
species, of which one, Jatifrons Carriker and Shull, is a Myrsidea. 

Ewing reviewed the genus in 1930 (Proc. U. S. Nat. Mus., vol. 77, 
art. 20) and listed eight valid species, of which two are Myrsidea 
(femuralis Kistiakowsky and piageti Kistiakowsky). He placed 
bruneri Carriker under the synonymy of dubius Kellogg, stating that 
Ferris had examined a cotype of it and pronounced it to be the same 
as dubius, in which he agreed; but this statement is clearly an error, as 
will be shown later in this paper. He also described three new species 
(richmondi, spiniger, and australis). On page 2 he says, “‘latifrons 
Carriker and Shull . . . is a true Dennyus,’”’ but he does not include 
it in his list of species of the genus. He redescribed and figured 
dubius Kellogg and, on the whole, left a much clearer picture of the 


genus than we previously had. 
284245—54 533 


534 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


In 1941 Thompson (Ann. Mag. Nat. Hist., ser. 11, vol. 7, p. 530, 
1941) described a species (francicus) from the Solomon Islands and 
in 1948 (Bol. Ent. Venezolana, vol. 7, Nos. 1, 2, June 1948) he de- 
scribed a species from Ceylon and another from Jamaica (cypsiurus 
and gossez). 

Because of the small number of specimens and species available 
for study to any single worker, it has been and still is impossible to 
prepare a thoroughly comprehensive review of the genus. I have been 
fortunate in securing a considerable number of species of this inter- 
esting group. In my collection are the types of brunert (Carriker) 
and meridionalis (Carriker), several specimens of australis Ewing from 
the type host, and a male of distinctus Ferris. 

In addition, there are specimens of seven forms which appear to be 
undescribed, all from various countries of South America. Some of 
these species are very distinct and may be easily recognized, but 
others form a closely related group, some of which may be conspecific 
but may be separated from each other by the shape of the head and 
thoracic segments, chaetotaxy, etc. I have examined males of seven 
species (four described species and three new ones) and found very 
little difference in the genitalia (see figures). Even the genitalia of 
distinctus Ferris, one of the most aberrant species of the genus, are 
very similar to the others. Apparently, the only very different 
genitalia are those of major (Uchida), for which Uchida erected a new 
genus not recognized by Hopkins and Clay. The males, insofar as I 
can determine, differ but little from the females except in size and in 
the terminal segments of the abdomen. 

The best characters for the separation of the species are the gular 
plate and its chaetotaxy, the shape and proportions of the thoracic 
segments, the prosternal plate and its chaetotaxy, the mesosternal 
and metasternal plates, and a median sternal sclerite usually present 
in abdominal segments I and II (sometimes wanting). 

The following key contains the new species described in this paper 
as well as all of the known species except cypsvurus Thompson, 
francicus Thompson, and hirundinidis (Linnaeus), the European 
species for which I lack sufficient data for their inclusion. I have 
followed Peters in the nomenclature of the hosts. All hosts whose 
identities were uncertain have been verified by Dr. A. Wetmore except 
that of D. similis, which was furnished by Mr. Phelps. To both of 
these my thanks are due. All measurements are in millimeters and 
all drawings were prepared by the author. 


NEOTROPICAL MALLOPHAGA—CARRIKER jae 


Key to the species of Dennyus 


(Data are insufficient for the inclusion of cypsturus Thompson, francicus Thomp- 
son, and hirundinidis (Linnaeus).) 


la. Head very short and wide; prothorax quadrilateral, without lateral lobes; 
mesothorax and metathorax very short and very wide (more than 3 times 
wider than long); abdomen oval_____--__-_-_--- brevicapitis, new species 
1b. Head but little wider than long. 
2a. Pre-antennary portion of head with sides almost parallel and front trans- 
verse (head, .49 by .51); temples more or less angular_-_distinctus Ferris 
2b. Pre-antennary area circular; temples small, narrow and rounded, but little 
Preanged qnesus47 Dy B92 2208 2 Se eee eS minor (Kellogg) 
lc. Head, thorax, and abdomen of shape normally found in the genus. 
3a. Fringe of setae along anterior edge of temples double, one pointing out- 
ward, the other backward; double rows of long hairs on posterior 
margins of temples, occiput, prothorax, and metathorax; patches of 4 
to 6 exceedingly long hairs in median portion at each side of tergites 
II to VII (nearly as long as abdomen) ______-__- intonsus, new species 
3b. Chaetotaxy normal, not as above. 
4a. Posterolateral margins of temples clearly convex from lateral to poste- 
rior angle, not straight or concave. 
5a. Abdominal segment VIII in both sexes deeply concave on poste- 
rior margin, with all (#) or most (9) of segment IX within this 
concavity; Male genitalia very different from all other species 
Bee peu Users tees seek ee ees ene major (Uchida) 
5b. Abdominal segments VIII and IX normal, not as in 5a. 
6a. Numerous peglike spines on dorsal surface of head and prothorax. 
spiniger Ewing 
66. No peglike spines on head or thorax. 
7a. Prosternal plate with numerous short, thickened spines (10 to 12), 
with or without longer setae. 
8a. Pre-antennary margin of head uniformly rounded and poste- 
rior margin of temples strongly convex; 10 short spines on 
prosternal plate, without any longer setae. 
rotundocapitis, new species 
8b. Pre-antennary margin of head not uniformly rounded, but with 
sides slightly concave. 
9a. Twelve short, straight spines and 2 long setae on prosternal 
plate; plate with straight sides converging sharply 
posteriorly; temples strongly convex posteriorly. 
brunneitorques, new species 
9b. Twelve short, curving spines on central area of prosternal plate, 
and a long, curving seta at anterior angle of same area; 
plate with sides subparallel and concave_-_richmondi Ewing 
7b. Prosternal plate with 3 to 6 longish setae on anterior portion. 
10a. Prosternal plate long, and narrow posteriorly, with narrow 
lateral bands, and with 5 (or 6) longish setae; temples 
strongly rounded and pre-antennary margin uniformly 
circular. 
lla. Setae along posterior margin of abdominal tergites consist 
of short, thickened hairs, alternating with hairs nearly 
length of segments; sides of thorax straight. 
similis, new species 


536 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


11b. Setae along posterior margin of abdominal tergites consist 
of slender hairs, sparsely set, and mostly longer than the 
succeeding segments. .__-----------.-- australis Ewing 
10b. Three longish setae on prosternal plate, one on each side on 
anterior border and one in anterior center of inner, clear 
area; temples very slightly convex and pre-antennary 
margin with slight concavity on each side__-gossei Thompson 
4b. Posterolateral margins of temples straight or concave (never convex). 
12a. Posterolateral margin of temples straight; occiput transverse; 
prosternal plate large, with slightly concave, converging sides and 
wide marginal bands; 11 or 12 short spines set on inner portion 
of plate and a long hair at anterior, lateral angles of inner portion. 
spininotus, new species 
12b. Posterolateral margins of temples decidedly concave just forward 
of the posterior angle. 
13a. Metathorax normal, with posterolateral angles simple and connected 
by a submarginal band to abdominal segment I (see fig. 68). 
14a. Prosternal plate with 3 to 5 setae; when only 3, all are long; 
when 5, one is very small and 2 medium. 
15a. Prosternal plate with 2 long and 2 rather short setae, all set 
in anterior marginal band of plate, with the 2 short ones 
set medially (often a very short spine at one side). 
limbus, new species 
15b. Prosternal plate with 3 long setae, set in anterior portion of 
medial clear area; temples small; head narrow. 
dubius (Kellogg) 
14b. Prosternal plate with about 8 longish setae, all set within the 
median, cleanspacesas 2] ae oa. ae eee bruneri (Carriker) 
136. Metathorax with posterior margin much wider than abdominal 
segment I, with the posterolateral angle bifurcated; the spines 
along the posterior margin of segment beginning at the inner 
point of the bifurcation, also connecting band across to the 
SDC Omen 232-2 cs aie eae en a alene meridionalis (Carriker) 


Dennyus australis Ewing 
FIGURE 63,7 


Dennyus australis Ewing, Proc. U. 8. Nat. Mus., vol. 77, art. 20, p. 5, 1930. 

(Host, Apus a. andecolus (d’Orbigny and La Fresnaye).) 

I have in my collection one adult male and two immature females 
from Apus andecolus parvulus (Berlepsch and Stolzmann) collected 
at Huarmey, Pert, and one adult female from the type host collected 
at Oploca, Bolivia. The type of australis is a female, and my female 
from Oploca agrees closely with Ewing’s description and figure of the 
species, although the measurements differ slightly, those given for 
the type being 2.75 by 1.30, while my female measures 2.86 by 1.19. 

The male from A. andecolus parvulus has exactly the same shape, 
structure, and chaetotaxy of the head as the female, as well as the 


NEOTROPICAL MALLOPHAGA—CARRIKER 537 


shape and chaetotaxy of the prosternal plate as given by Ewing. It 
measures 2.36 by .90. 

This species is close to bruneri (Carriker), having the same style 
of head bands and exactly the same gular plate and chaetotaxy. It 
differs, however, in the shape of the head, australis having the temples 
convex, while in brunert they are concave. In australis the sides of 
the mesothorax and metathorax are straight, while in bruneri the sides 
of the mesothorax are convex and the sides of the metathorax are 
concave. JD. australis is also much larger than bruneri, the male of 
the latter measuring only 2.13 by .80. The dorsal aspect of the 
paratergals is different, as well as the shape and chaetotaxy of the 
prosternal plate, that of bruneri being shorter, and broader anteriorly, 
and with the eight longish hairs all set inside the clear area, while in 
australis there are four setae in the clear area and two in the lateral 
band (male). Ewing shows three setae in the lateral band and three 
in the clear area (female), but in my female the setae are arranged in 
the same manner as in the male. 

Measurements of adult male and female in collection of author: 


oh Male a Bema 
Length Width Length Width 
Ee eee Ake Se ee ee ROO ge oe yee COTE ee 
SUES Ot a a kt pe te ie ok BR mea ipa tS Reg ah ca oA ee pen a ae” SENT SA 
SG RORSACE asus LeU FFt5T Sh) {tle RIE Le AGM Ad paah a ve OL 
eeeerrs: ee ee serocis eam. 054 a . 49 . 65 . 542 . 76 
ee Se eres wee eee ne Aan pe be 32 sBiiy Jsacals 
ec hOLaKe spe ve se ee Ok Bae . 288 . 40 . 314 . 465 
Mesa-metathorax. 202200. 22028 2 oto pti 655 37 803 
PDH UTneTnEaere ice re ce Ee AA! 1, 44 90 1. 86 1.19 
Penremapinge, ers Abn. EL FE od BU .41 Bt OAL SOIAOTY Glee . 230d 
Paesimneres er an maw) ayes’) oy hye . 24 SI4iy myri san fads 
Hndomeral’ 8305. 326 ee 2's. ons oe cease 217 1 ns cett yh eG lee 


Dennyus dubius (Kellogg) 


Nitzschia dubius Kellogg, Proc. California Acad. Sci., ser. 2, vol. 6, p. 540, 1896. 

(Host, Chaetura pelagica (Linnaeus) .) 

I have no specimens of this species and have used, with some 
misgivings, Ewing’s description and figures as a basis for my con- 
clusions regarding it. Ewing says his description is based “upon 
many adult specimens chiefly from the type host’’ (italics mine), and 
lists under dubius specimens not only from Aecronautes melanoleucus 
(A. s. saxatilis), type host for D. brunert (Carriker), but from Chaetura 
richmondi, from Eden, Nicaragua; yet, in the same paper, he describes 
Dennyus richmondi whose host is Chaetura richmondi, also from Eden, 
Nicaragua. 


538 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


I have in my collection two males (paratypes) of D. bruneri, from 
Aeronautes s. saxatilis, and they certainly are not what Ewing describes 
as dubwus (see key). Although the two are undoubtedly closely 
related, and may possibly be conspecific along with the new 
species (limbus) described on page 542, until more is known about 
this group I prefer to leave them as separate species. 


Dennyus bruneri (Carriker) 


FicurE 63,c 


Nitzschia pulicaris var. tibialis Carriker, Journ. New York Ent. Soc., vol. 10, 
p. 225, pl. 22, figs. 4, 5, 1902. (Host, Aeronautes melanoleucus (=A. s. saxatilis 
(Woodhouse).) 

Nitzschia bruneri Carriker, Univ. Nebraska Stud., vol. 3, No. 2, p. 55, 1903 
(new name for N. tibialis, preoccupied by WN. tibialis Piaget, 1880). 

I have in my collection two males (labeled as types) of this species. 
The original type series contained three males and one female. I 
imagine that one male and the female remained in the collection of 
the University of Nebraska, and if they still exist they should be 
considered as the male holotype and female allotype of the species. 

This species may be distinguished from its closely related forms 
by the detailed characters as given in the key. From dubius it may 
easily be distinguished by the chaetotaxy of the prosternal plate, 
dubius having three longish setae set in a row along the edge of the 
clear area, while in bruneri there are eight longish setae irregularly 
placed within the clear area. Ewing’s figure shows the posterior 
margin of the prothorax with sides angulated and median portion 
strongly concave, while in bruneri the posterolateral angles are rounded 
and the posterior margin is almost transverse. In dubius the meta- 
thorax has the sides straight and is ‘‘over twice as broad as long,” 
while in bruneri the sides of metathorax are concave and its width 
is three times the length. Ewing’s description of the abdominal 
chaetotaxy is very vague but I suspect that it is different from that 
of bruneri, which has one spine in lateral angle of segment I and one 
slender, submarginal hair; in segment II there are one spine and one 
slender hair in the angle and a longer, coarser, submarginal dorsal hair. 
In segments III to VII there are two short hairs in the angle and 
one long submarginal hair, with two very long hairs at angles of 
segments VIII and IX. These long dorsal hairs become progressively 
longer posteriorly. 

In dubius the parameres are incurved apically, their tips touching, 
while in bruneri they are almost straight in one specimen and slightly 
incurved apically in the other. 


NEOTROPICAL MALLOPHAGA—CARRIKER 539 


Measurements of the male paratype: 


aetrt fee ee ee Sees Dias 23S 
feces eee ee re ed ee ee ee eee ee. LS 
Tb TORSAG wee ks = i eee a a bo te eee 456 
URED T Ett 2a 2 ee Eee oS a es . 48 . 63 
TOPEY ETERS. ewes Agi 2 LS re “S| ee AAR s Tas 
prop Roraxee ee en So ot ee em ee eae . 26 . 38 
Mesoumetathoraxe. 20 Se oe ee oy ee . 326 . 615 
Pipa Grn nieeseee mete ee. SAP OE 2 eee es 1. 26 . 80 
REeREMETCrbederaserys) Sey) Re 2 ef OB .14 


Dennyus meridionalis (Carriker) 
FiGcureE 63,6,h 


Nitzschia bruneri ‘var. 'meridionalis Carriker, Univ. Nebraska Stud., vol. 3, 
No. 2, p. 56, 1903. (Host, Chaetura griseiventris (=C. spinicauda fumosa 
Salvin).)! 


I have in my collection a slide containing a male and a female of 
this species, labeled as ‘‘types,’’ and another slide, also containing a 
pair of adults and several nymphs, labeled merely ‘“‘Pozo Azul, Costa 
Rica, June, 1902.” 

Eight skins of the host swift were collected by me at Pozo Azul, 
and all are in the Carnegie Museum. It is probable that the various 
specimens of Dennyus came from more than one of these birds. This 
fact is mentioned because of the presence of two apparently distinct 
species in this series, including specimens from Chaetura spinicauda 
aetherodroma Wetmore, from Colombia. The whole series seems to 
be practically the same in every way except in the structure of the 
metathorax, of which there are two very distinct types (see figs.). 
It does not seem possible that this can be a case of individual varia- 
tion, since the difference is very great and clear-cut. 

The male and female labeled as the types of meridionalis are of one 
class, while the male and female ‘‘paratypes” are of the other. A 
female from Cartagena, from OC. s. aetherodroma Wetmore, has the 
metathorax as in the “paratype’’ of meridionalis, as have also a female 
taken at El Real, Colombia, from the same host and a female taken 
at Las Vegas, Colombia; but a male from El Real is like the types of 
meridionalis. 

All the above specimens have the chaetotaxy of the prosternal 
plate the same except the female from Las Vegas, which has only 
three rather short, spinelike setae in a straight row across the anterior 
marginal band, while the plate itself is longer and narrower than in 

1 Hopkins and Clay (A Checklist of the Genera and Species of Mallophaga, 1952) have given the host of 
this species as Chaetura griseiventris phaeopygus Hellmayr, a bird found on the Caribbean side of Costa 


Rica. The birds from Pozo Azul (Pacific side) were identified by Todd as well as by Wetmore as 
Chaetura spinicauda fumosa Salvin. 


540 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


ceed § 


; 
| 


Ficurs 63.—For explanation see facing page. 


NEOTROPICAL MALLOPHAGA—CARRIKER 541 


Ficure 64.—a, Dennyus spininotus, new species, female; b, female genitalia of D. rotwndo- 
capitis, new species; c, D. brunneitorques, new species, female; d, antennae of D. rotun- 
docapitis, male; e, D. similis, new species, temale (patches of sternal setae to right of 
figure); f, D. rotundocapitis, female; g, male genitalia of D. similis. 


Ficure 63.—a, Dennyus limbus, new species, female; b, type, female, of D. meridionalis 
(Carriker); c, paratype, male, of D. bruneri (Carriker); d, thorax and abdominal segment 
I of female D. limbus; ¢, D. intonsus, new species, female; f, male genitalia of D. limbus; 
g, D. brevicapitis, new species, female; h, thorax and abdominal] segment I of D. merid- 
ionalis; i, male genitalia of D. australis Ewing. 


284245542 


542 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 108 


the other specimens. It is possible that the Las Vegas (in Sierra 
Nevada de Santa Marta) birds are not typical C. s. aetherodroma. 
Todd originally called them C. s. spinicauda, saying that they were 
like birds from Venezuela and Trinidad, although Dr. Wetmore lists 
the Las Vegas birds as that race in his description of aetherodroma. 

The only solution to this problem seems to be the separation of the 
series into two species, the male and female types of meridionalis 
and the single male from El Real comprising one species (meridionalis), 
while the pair of ‘“‘paratypes” of meridionalis and the three females 
from Colombia form the other, which I have described below as 
D. limbus, new species. 

D. meridionalis may be recognized at a glance by the peculiar 
structure of the metathorax in conjunction with the prosternal plate 
and its chaetotaxy. 

Measurements of the types: 


Male Female 
Length Width Length Width 
Body eee es ot Eo ree eee 701 Pe eee 2. (1) 2 ee 
Head 
ant Hossget were. Je oe ee eee 477. eee . 50 
Pr S1L Cero oy dap idate (ty SN eee ae aN A =o . 694 . 545 . 745 
OCCU se Ua gee 2 an OE — Ae ee 2456) (Soe ATA See 
Prothoraxe tek ae. a ees, Se eae: . 29 41 1326 . 435 
Meso-metathoraxs:.. -. 2°) Sebi ae 38 814 435 976 
Abdoment=< sue Ser (2! Ags. a ee 1. 20 7057 te? 1. 02 
Paramenesiqs 2 Oi: 24 SNe so ener me 206 138) 4 See 


Dennyus limbus, new species 
FicureE 63,a,d,f 


Types.—Male and female adults from Chaetura spinicauda fumosa 
Salvin, collected by the author at Pozo Azul, Costa Rica, June 1902 
(in collection of author). 

Diagnosis.—Closely related to dubius (Kellogg) and bruneri (Car- 
riker), having the same concave posterolateral margins of the temples 
and the same thoracic structure. 

D. limbus differs from bruneri in the wider head (.49 by .684 against 
A8 by .63) and much wider thorax (.76 against .615); the abdomen 
(females) is smaller (1.20 by .716 against 1.26 by .80). It differs 
from both dubius and bruneri in the shape and chaetotaxy of the 
prosternal plate. The plate is much larger than that of bruneri, 
having four setae (two long and two short) set in the marginal band, 
while bruneri has eight shorter setae, all within the clear median area. 
D. dubius has but three long setae, set at head of median clear area. 
A discussion of the metathoracic characters is given under remarks on 
meridionalis and in the key. 


NEOTROPICAL MALLOPHAGA—CARRIKER 543 


Measurements of the types: 


Male Female 
Length Width Length Width 
ME ae a ek DOT (hse eek FT piel a telah te 
Head 
RETO BLOSSRO Ce oe es Ne es ARE SAGO . 51 
emer oss ete ee Bi iet fees . 49 . 684 . 542 . 755 
ACC LES Lees y= Yeeeah ee Doky! eae Soper re Rohe Bee 
REECE RUIGSI CER ME ete ee ee ol oy Me eS . 28 . 40 5 BvAS . 467 
Wrese-metauhorax.. §0 9. 2 Pie egy oak _ Bs 76 . 42 . 74 
mNdaMoner 2 ee ek eh 1. 20 WGI s [68s 1. 04 
“ACE G eis ed le ee oe ene OS ee _ 220 rit ps PO Spd ie ee nt et 


Dennyus gossei Thompson 


Dennyus gosset Thompson, Bol. Ent. Venezolana, vol. 3, Nos. 1, 2, p. 1, June 

1948. (Host, Tachornis p. phoenicobia Gosse.) 

This species is well described and figured by the author and needs 
little comment beyond the characters given in the key. The occipital 
margin is deeply reentering, with occiput strongly convex; the shape 
of the gular plate also seems to be characteristic; the temples are 
very flatly convex and the prosternal plate characteristic with its 
straight, converging sides, narrow borders, and three very long setae 
(almost as long as plate), two in marginal band and one in clear area. 


Dennyus distinctus Ferris 


Dennyus distinctus Ferris, Canadian Ent., vol. 48, p. 310, 1916. (Host, Callocalia 
sp. from Java.) 

A single male of this interesting species was collected by R. H. Baker 
from Callocalia inexpectata bartschi Mearns on the Island of Guam on 
July 29, 1945. This specimen was previously reported on and figured 
by the author (Proc. U. S. Nat. Mus., vol. 100, p. 16, 1949). 

D. distinctus is probably the most abnormal species of the genus as 
regards the shape of the head, which is about as long as wide, with a 
wide, transverse frons and sides of pre-antennary area straight and 
slightly divergent. The meso-metathoracic suture is faintly visible, 
but the two segments are very closely united (this detail omitted in 
my figure in paper cited above). 


Dennyus major (Uchida) 


Takamatsuia major Uchida, Journ. Coll. Agric. Tokyo, vol. 9, p. 32, 1926. (Host, 

Hirundapis c. caudacutus (Latham).) 

A new genus was erected by Uchida for this species on the strength 
of the shape of abdominal segments VIII and IX in both sexes and 
the male genitalia. Segment IX is very small (especially in the male) 
and enclosed within a deep emargination of the posterior margin of 


544 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Segment VIII. The male genitalia is also quite different from all 
others of the genus which I have seen, the remainder being unusually 
uniform. I have carefully examined Uchida’s description and figures 
and am of the opinion that this species may be entitled to subgeneric 
rank, but without actual examination of specimens I hesitate to 
do this. 


Dennyus minor (Kellogg and Paine) 


Nitzschia minor Kellogg and Paine, Rec. Indian Mus., vol. 10, p. 243, pl. 15, 
fig. 10, 1914. (Host, Apis a. affinis (J. E. Gray).) 

Like most of Kellogg’s descriptions, this one is very inadequate and 
the figure not particularly illuminating. The species seems to be 
characterized by its small size, circular frontal margin of head, and 
small, rounded temples. He gives the following measurements: 


Male Female 
Length Width Length Width 
ody 2 8 2 Le AES AES Ee A lh Toe Eee hy D2 ee 
Heade 59-0 Fa aio ole patio tapesnays . 39 52 47 59 
IPTOtHOTAKASS 28 8 ee ae A ee . 22 32 23 36 
Metathorascts 280 5 25 ae ee eo 20 48 32 58 
VeN OG CORGOYE) 1s, = 2 Oe me ee a Pet a . 87 71 90 71 


Dennyus brevicapitis, new species 


FIGureE 63, g 


Type.—Female adult from Chaetura b. brachyura (Jardine), collected 
by the author at Carenage, Trinidad, B. W. I., August 14, 1909 (in 
collection of author). 

I bave been undecided as to the generic position of this species, 
whether it belongs in Eurewm or Dennyus. Certainly it presents 
characters common to both, resembling the former genus in its wide, 
short head, rectangular prothorax, and oval abdomen. It might be 
classed as the connecting link between Dennyus and Hureum, but on 
the whole it seems best to place it under Dennyus. 

Diagnosis.—The species is so different from all other known species 
of the genus that, in addition to the figure given, very little description 
seems necessary. 

The head is nearly twice as wide as long, with pre-antennary area 
flatly conical, and sides flatly convex; temples short and expanded 
laterally; prothorax quadrilateral in shape, encircled by a wide, deeply 
chitinized band, with three spines on anterior angles, two long hairs 
on the rounded posterior angles and two on posterior margin. The 
prosternum is of the usual type for the genus, witb four short spines 
and two longer, thickened setae, all set in the marginal band. 


NEOTROPICAL MALLOPHAGA—CARRIKER 545 


The mesothorax is very short and wide, with acute lateral angles 
and strongly convex posterior margin. The metathorax is nearly 
four times as wide as long, with posterior margin flatly poited medi- 
ally. The greater portion of the paratergal plates lies on the ventral 
side; they are wide, heavily chitinized, and deeply colored, with 6 to 
12 spines on posterior margin; the dorsal portion of pleurites II to 
VIII contains heavily chitinized incrassations of varying shapes, no 
two being exactly the same. 

The legs are normal for the genus, but the third pair of femora are 
unusually short, scarcely longer than the second pair. The chaetotaxy 
of the head and mesosternum is typical of the genus. The long hairs 
at the lateral angles of tergites II to VIII are thick and long, there 
being one or two in each angle which are thicker and longer than the 
others. The dorsal chaetotaxy of tergites IT, III, and V is not shown 
in the figure, but is approximately equal to that of IV and VI, the 
hairs becoming progressively longer posteriorly, with one or two 
shorter hairs between the longer ones which reach across the succeed- 
ing segment. The female holotype is the only specimen of the species. 

Measurements of the type: 

Length Width 


Dien yemerrtesis Of ead teh Isis, SOILS VAULT Oe  e 2n21i6 Swale 
to arlene epee ay a he pays ets ee ae Ld ie ical Bie By alle eg aa fag se 
DON, LENSER (eg Si aes SOOT ey Ate ame oe ee Bee, SOR OYE Sear Serre ee See ae ae . 52 
PRES. _ See ee OR ee ee eee . 456 . 80 
ACTON fie wd erage “paces ee Seat Se oe raed Ee Mh i eer me On Romeo 
TOD OLsaees ee IN a A eee . 303 . 467 
enm-Ine pa biOraxteise Sie -ST0y Cage RATE Mae ee Pere Ae . 38 . 88 
Pape Ae She koh oer cd ole eet ee SO Le 1. 36 1. 10 


Dennyus intonsus, new species 


FIGURE 63,e 


Type.—Adult female from Chaetura chapman viridipennis Cherrie, 
collected by the author at El Real, Rio Cauca, Colombia, March 8, 
1948 (in USNM). 

Diagnosis. —D. intonsus is a very distinct species and may be 
recognized at a glance by the abnormal chaetotaxy of the whole body. 
There is a double fringe of setae along the anterior margin of the 
temples; one fringe is along the edge, pointing outward, and the other 
is submarginal, parallel to the first, and pointing inward and back- 
ward. There are two rows of very long hairs along the posterior 
margin of the temples, the usual four along the edge and an additional 
row of five submarginal; also four long, dorsal hairs on the occiput, 
two near the margin and two others, longer, inside the marginal pair; 


546 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


There are three spines on the anterolateral angle of the prothorax, 
and two more posterior to them; the three long marginal hairs on 
posterior edge of prothorax are also duplicated by three longer ones 
just inside them. There are eight hairs (four long, four short) on each 
side of the posterior margin of the metathorax with an additional 
submarginal row of nine much longer ones. 

The chaetotaxy of the abdomen is exceedingly complex, but may be 
understood from a careful study of the figure. However, it was not 
possible to put in the figure all of the hairs on all of the segments. 
The most striking feature of the abdominal chaetotaxy is the groups 
of six extremely long hairs set in the median portion of tergites II to 
VI, just within the lateral margins. These hairs are almost as long as 
the abdomen. In the same position on tergite I there are three long 
hairs and a spine, while on tergites VII to [X there are two long 
submarginal hairs in addition to the other long ones at the angle. 

The first and second pairs of legs are normal, but the third femur is 
short and slender, scarcely longer than the tibia. The setae forming 
the patches on the third femora and abdominal sternites are very fine 
and closely set, those on the abdomen forming a continuous patch 
from middle of sternite V to near end of sternite VII. 

The structure of abdominal pleurites and sternites is unusual and 
not clearly determined, the posterior margin of the sternites not 
reaching to the lateral edges of the abdomen. The head is unusually 
small and the thorax narrow. The type apparently is an adult female 
recently moulted so that all of the chitinized bands are not clearly 
outlined. The characteristic sternal plate on segments I and II is 
apparently absent. The prosternal plate is normal and contains four 
long and two short, slender setae. 

In addition to the type there is a nymph which has a quite different 
chaetotaxy, having not yet acquired the duplicated temporal fringe 
and the patches of long hairs on the tergites (only one long hair and a 
spine on tergites II to V, and two long hairs on VI to IX). It is 
impossible to determine with certainty the sex of this nymph, but it 
seems, from the structure of abdominal segments VIII and 1X, to 
be a female. 

Measurements of the type: 


Length Width 


Body ec it tet Mat Beh eis eh I RIT OD Oe 2h he PERO. Sa SSS Ae 2: 204 Sees 
PU a Lehi Sec hak Se a Ne) REO A a 
ants fossac=-. bets eee ult see dle eset base eee . 456 
tem plese is! ices Se Pe ee . 47 . 655 
OCCIPUG 25 ne Lg Cia cen tate teat 2 RU ae Ree ight = ae  Ad4 | cee 
Prothorax See ee ee ee ee ee eee . 267 . 282 
Meso-smetathordxe. oe 222 0 eR ean ee eee) ee 355 654 


NEOTROPICAL MALLOPHAGA—CARRIKER 547 


Dennyus spininotus, new species 


Ficure 64,a 


Type—Female adult from Cypseloides fumigatus (Streubel) 
collected by the author on Rio Esmeralda, Department of Cérdoba, 
Colombia, May 19, 1949 (in USNM). 

Diagnosis.—The general shape of the head resembles very much 
that of D. gossei Thompson, although the temples are perfectly 
straight on posterior margin, but the measurements of the body are 
very different, 2.80 by 1.02 against 2.30 by .81 for gossev. 

Unfortunately the type, and only specimen, is in poor condition, 
having been cleared too much and having collected within the head 
and thorax a quantity of food particles which obscure the few visible 
details of internal structure. 

The whole thorax and abdomen are of normal shape and structure, 
presenting no outstanding characters. The chaetotaxy of the head 
is also normal, except for the three dorsal spines which are present in 
but few species. The prosternal plate is unusually large, very broad 
anteriorly, and with wide marginal band. There is a long, slender 
seta at each anterior corner of the clear area and 11 heavy, short 
spines irregularly set within it. There are numerous heavy spines 
on the mesosternum, those beneath the posterior portion of the 
prothorax being thicker than the patch on the posterior margin of the 
mesosternum. 

The spines along posterior margin of the tergites are thick and 
closely set, interspaced irregularly with long setae, usually 2 to 3 
spines between the long hairs: there is but one longish hair at the 
posterolateral angle of tergites II to VII. Short, thick setae are 
scattered irregularly over the sternites, while the patches of setae on 
sternites V and VI are sparse and coarse, as well as those on the third 
femora; there are an unusual number of short, thick spines along the 
outer edge of the second and third femora, and the spines along pos- 
terior edge of ventral pleurites are unusually numerous and thickened. 
The peculiar median sternal plate usually present on segment I of 
abdomen seems to be absent, at least it is not visible. 

Measurements of type: 


Length Width 


Eee SEI YL heels (Due ee ee ettyy So e ag sr 2E0y nssiass 
es een es thd a ren Byer ae De pe Rd oat yed is 2 weppte ey) oes 
STIR ARLORS 2G ee hs RW Ne eB es Ee pa NS a 5 eo 586 
PTL TT 2. oes at Ra i lS age eR a . 521 . 836 
LIVE eget yest ib el leper teppei > gic eat BE een Alar 146" oot eee 
PENG GONAI eee eee! SHES EEE SEEN) ASE AML OPS SAL Shah. s8V/ . 471 
Niesa=metathoraxisriey. ait) _ tay SOR i Sib te mer eee + Oi 91 


IN WoVevae(e\olse been Sa LAT OP Oe cs Eee eee ents? Oy Bae Wg Mee ty ete me 1. 65 1. 02 


548 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Dennyus rotundocapitis, new species 


Fieure 64,6,d,f 


Types.—Male holotype from Streptoprocne zonaris albicincta 
(Cabanis) collected by the author near Cali, Colombia, December 16, 
1950 Gn USNM), and female allotype from same host collected by the 
author at Huancano, Pert, March 9, 1931 (in collection of author). 

Diagnosis.—This species may be recognized by the narrow, broadly 
rounded temples, without angles, the flatly rounded pre-antennary 
margin, the prosternal plate and setae, the sparse, coarse setae of 
femoral and abdominal patches, and by the shape of the sternal plate 
on segments I and II. 

The bands of chitin which connect the prothorax with the meso- 
thorax and the metathorax with the abdomen are unusually heavy, as 
are the dorsal incrassations of the paratergal plates. The prosternum 
contains 10 short, thick spines, all within the clear area. The species 
is represented only by the male holotype and the female allotype. 

Measurements of the types: 


Male Female 

Length Width Length Width 
Bo@ivea. seead. 7: pear ce pee ezad’T <2 2: O04) pur _4a2. 2 2: G31 | eee 
Pea a a eee es eters eho Ea ten Ne Src Ek cassie 
Rnb LORSAGE foto oe a eee ee re ey Bae _ 49° ae 564 

temples? ssf ee Se ee ee ne ee . 46 . 673 . 55 er ied 
OCCIPUtY £448 Sh etee Marre: Sot eae NASA pee: ae ASh (eee 

Prothorasi> 2 297 00k at. ine gin . 303 . 39 sate . 48 
Meso-metathorax__.____.______-_-_____-_ . 825 62 445 835 

PBUONIBIAA 0 eco 8 ol eee 1. 16 78 1. 67 1.11 
PATHINGLER ooo ee eee ee ae . 228 13. eee 


Dennyus brunneitorques, new species 


Ficure 64,¢ 


Type—Female adult from Chaetura rutila brunneitorques La Fres- 
naye, collected by the author at San Juan, Chanchamayo, Pert, 
June 6, 1930 (in collection of author). 

& Diagnosis —Somewhat similar to rotundocapitis, new species, but con- 
siderably smaller (head, .467 by .694 against .55 by .77), with temples 
less rounded and more angulated both front and rear; pre-antennary 
margin with pronounced concavities on each side; occipital bands and 
gular plate very similar to those of rotundocapitis, as well as entire 
thoracic structure. The prosternal plate is of decidedly different 
shape and chaetotaxy, having three spines and two long setae along 
the border of the anterior band, and with nine spines set within the 
clear area. The shape and chaetotaxy of the terminal abdominal 
segments are also somewhat different. The median sternal plate on 


NEOTROPICAL MALLOPHAGA—CARRIKER 549 


segments I and II is absent. The abdominal chaetotaxy and lateral 
tergal incrassations are very similar to those of rotundocapitis. 


Measurements of type: 
Length Width 


I Eo eS eS eye is eos ane Zale, see a 
cigs ey ee ce es Bee aed Re eS ee ee eae See 
EOE U Re LACSEA BS aD els A a ee ee ee SET Sia By 
REINER oe he ete ee Suet i eee . 467 . 694 
LS Ure ae te ee oe ee ae Sa eee Ae ss ees 
PETES UGE Eg TS OR ca Pi ea a pe dg ete Brest . 412 
LMU ESY Pct ENED COS oS pean SE ES ne eel . 39 . 716 
Peat FIThIG ieee eee ie he ds eee ee eee eae ee aeee 1. 34 . 94 


Dennyus similis, new species 


Ficure 64,e,9 


Types.—Male and female adults from Progne c. chalybea (Gmelin)! 
collected by J. Pablo Anduze at Barinas, Venezuela, January 14, 
1943 (in Department of Hygiene, Caracas, Venezuela). 

Diagnosis —Somewhat similar to D. australis Ewing, from which it 
differs as follows: Gular plate differently shaped; temples more convex 
posteriorly; lateral wings of prothorax less pronounced and posterior 
margin slightly concave instead of convex; the sides of both mesothorax 
and metathorax are quite straight (except for slight swelling near suture 
on metathorax), while these segments in australis are convex on sides. 

The setae along posterior margin of abdominal tergites consist of 
short, thickened hairs, alternating with hairs nearly the length of suc- 
ceeding segments, while in australis the chaetotaxy consists of slender 
hairs, sparsely set, and mostly longer than the succeeding segments; 
the patches of setae on femora and abdominal sternites are composed 
of finer hairs, set closer together. The male genitalia also differs 
slightly, the parameres being shorter and endomeral sac longer, while 
the shape of the basal portion of the basal plate is quite different. 

Measurements of types: 


Male Female 

Length Width Length Width 
CPt. ~ hae BO eee ee POM See Qo) poe 
fetateal is Beye as oe eee renner eee gk - Se Ne ane ee been Se 

BRTET ROSS SUC eee et a ee reer SAGAS es Se 5 bY 
Semen es Soe ee .51 . 66 . 542 . 738 
Rees hei Se eee SADOM Cae = 40%) 
ES IEU IT G TS Kye ee ee aS . 25 . 369 . 303 . 423 

NexO-MEeLALNOIAX. = ok eo ee . 303 . 586 . 367 73 

Bana CA HENAOS YN eee 8 ae ae ee 1. 08 78 aaa 1. 01 
LET IPETETN ES ea net ee ea Pile 1 Dp Maines Mus ea es 8 


1 A note on the identity of this host is in press (1954) in the Boletin de Entomologia Venezolana. 


U. S. GOVERNMENT PRINTING OFFICE: 1954 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington : 1954 No. 3332 


NEARCTIC FLIES OF THE FAMILY PERISCELIDAE 
(DIPTERA) AND CERTAIN ANTHOMYZIDAE 
REFERRED TO THE FAMILY 


By A. H. Sturtevant ' 


The family Periscelidae was not recognized by earlier authors, its 
species being assigned to varying ones of the other acalypterate 
families. Even when it is recognized as a distinct group there is no 
agreement as to what genera should be included. My own studies 
have convinced me that the family is best limited (as concerns the 
Nearctic region) to the single genus Periscelis. (Nearctic species 
have been assigned to the genera Microperiscelis and Sphyroperiscelis, 
not here recognized as distinct.) 

There are included here a few notes on exotic Periscelidae, with a 
list (including references) of the known species of the world and a key 
to the known genera. I have included Cyamops since it has been 
included in recent accounts. I have also included Stenomicra, which 
has been referred here but neglected by some recent students. In order 
to place these two genera I have added a discussion of the Anthomy- 
zidae, with a key to the Nearctic genera of that family. 

The following abbreviations have been used in designating locations 
of type specimens: AMNH, for American Museum of Natural History; 
and USNM, for U. S. National Museum. 


1 California Institute of Technology, Pasadena, Calif. 
290839—54- 551 


002 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Family PERISCELIDAE 


Auxiliary vein rudimentary, ending free (not apically fused with 
costa or first vein); no costal breaks; anal cell present; first vein 
ending well distal to basal third of wing; costa ending at apex of third 
vein; second antennal segment with one or more dorsal apical bristles, 
rather long third segment arising from its ventral surface; arista 
plumose; face strongly receding below, extending laterally below the 
buccae, narrowed below the antennae by the anterior extension of the 
eyes (the longest diameter of the latter being oblique) ; bucca extending 
upward anteriorly, bearing a row of bristles, of which the anterior is 
placed well above the oral margin; hairs on lateral portions of face; 
one reclinate orbital bristle; no preapicals on tibiae; postverticals 
divergent; no presutural dorsocentral; scutellum with four marginal 
bristles, its disc bare. 


Genus Periscelis Loew, 1858 


I have suggested elsewhere that Microperiscelis Oldenberg (type, 
Notiphila annulata Fallén) is antedated by the isogenotypic Myodris 
Lioy and that the genus is scarcely valid. The properties of the new 
species described here have led me to conclude that Sphyroperiscelis 
Sturtevant is also best considered a synonym of Periscelis. There are 
11 or 12 species of the family known from the world, and five genera 
are recognized here, even though Myodris and Sphyroperiscelis are not 
accepted (see list, p. 556). For convenience, the use of broad 
generic limits is desirable here. If both the names under considera- 
tion are retained it will not be easy to fit the new species into any of 
the available categories, and a new genus will be indicated, giving 8 
genera for 12 species. 


Key to the Nearctic species of Periscelis 


1. Mesonotum dull gray, with a brown median stripe; posterior crossvein 
strongyisov «ud ) Jo. eobsede-g wes! alt do. feespa= P. annulata (Fallén) 
Mesonotum rather shining black; posterior crossvein faint or largely absent-_-2 

2. Antennal bases about as far from each other as each is from eye; posterior 


crossvein represented by stumps at each end____P. wheeleri (Sturtevant) 
Antennal bases much nearer each other than to eyes; posterior crossvein 
eompleéte but faints.o20 0. 0. bOI AY aS J 408 P. occidentalis, new species 


Female genitalia: I have described the internal reproductive 
systems of Periscelis annulata and P. wheeleri (Sturtevant, 1926, p. 7). 
These species are much alike, and are very different from any other 
known forms, especially in that they have three chitinized sperma- 
thecae that are attached directly to a singly common duct. They are 
also unusual in that the eggs are blackish brown in color. 


NEARCTIC PERISCELIDAE—STURTEVANT 553 


The European species of Periscelis are known to be most often 
found about the sap of bleeding trees (see, for example, Oldenberg, 
1914), and the South American P. nebulosa was also taken on a bleed- 
ing tree. All three of the Nearctic species are to be found in the same 
situations—most often on oak trees east of the Mississippi River, on 
cottonwoods west of it. I have reared P. wheeleri from larvae found 
in fermenting oak sap, and there can be little doubt that sap is the 
normal breeding place of all three Nearctic species. 

P. annulata has been taken in eastern Massachusetts from June 13 
until the end of August, in Missouri in June and July, in South Dakota 
in June, in Nebraska, New Mexico, and Washington in August, in 
Texas in October, and in Alabama on May 2. P. wheeleri has been 
taken in Massachusetts from June 18 to August 24; in New Jersey on 
July 3; and in California on October 24. P. occidentalis has been 
found in California from March 25 to July 30, in Arizona in June and 
July, and in Washington in August. Evidently all three species have 
numerous broods, probably overlapping, and may be found through- 
out the warmer months. 


Periscelis annulata (Fallén), 1813 


Described from Europe; recorded by Sturtevant (1923) from Ala- 
bama, South Dakota, and New Mexico, and by Malloch (1915) from 
Illinois. From the following locations 48 specimens were examined: 
Budapest, Hungary (Kertesz); Falmouth, East Falmouth, Woods 
Hole, and Naushon Island, Mass.; Webster Grove, Mo. (H. D. 
Stalker) ; Kushla (Mobile County), Ala.; Austin, Tex. (M. R. Wheeler) ; 
Chadron and Oakdale, Nebr. (M. R. Wheeler); Chamberlain, S. Dak.; 
Mogollon, N. Mex. (M. R. Wheeler). Dr. Wheeler informs me that 
he also has collected this species at Verlot, Wash. The specimen from 
Mogollon has the wings clouded on the anterior margin and it may 
represent a distinct form. 

The species was described in the ephydrid genus Notiphila and often 
has been placed in the Drosophilidae; I formerly referred it to the 
Agromyzidae, and later to the Lonchaeidae; Malloch (1915) has listed 
it under the Sapromyzidae. This species is the genotype of Myodris 
Lioy, of Microperiscelis Oldenberg, and of Meronychina Enderlein 
(Meronychia Enderlein). It was described by Malloch (1915) as 
Phorticoides flinti (genus and species both to be included in the 


synonymy). 
Periscelis wheeleri (Sturtevant), 1923 
Type from Naushon Island, Mass. (AMNH), and a paratype from 


the same locality (USNM). From the following locations 24 speci- 
mens were examined: Woods Hole and Naushon Island, Mass.; 


554 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Mendham, N.J.; and Pasadena, Calif. (M. R. Wheeler). The Cali- 
fornia specimen is most unexpected, but it is certain that it was col- 
lected in Pasadena, and it closely resembles eastern specimens. 

This species is the genotype of Sphyroperiscelis Sturtevant. 

The puparium is rather flattened, tapering at both ends, and, not 
including posterior spiracular processes, is nearly three times as long 
as broad. Surface dark brown, granular. There is a conspicuous 
thornlike process on each lateral margin of each abdominal segment. 
The posterior spiracles are on tapering divergent processes, each of 
which is nearly half as long as the maximum breadth of the puparium. 
There are no stalks to the anterior spiracles. 


Periscelis occidentalis, new species 


Male. Antennae yellow, brownish dorsally. Frons dark brown, 
subshining. Face pale yellowish, brown below centrally; entire oral 
margin silvery white. Mesonotum, scutellum, and humeri sub- 
shining black. Pleurae brown. Legs pale yellow, two dark brown 
annular bands on each femur and each tibia. Wing with rather in- 
distinct broad median darkened area (including anterior crossvein, 
most of first vein, basal section of third vein, and two basal sections of 
fourth), and a dark tip. Abdomen black, subshining. 

Arista with about 5 rather short branches above, 3 below. No 
differentiated vibrissae. Face in profile concave above oral margin, 
with a hump below antennal insertions. Eyes oblique. Frons about 
three-fifths width of head, as seen from above. Postverticals strong, 
divergent. Two pairs of dorsocentrals, both postsutural. Acrostichal 
hairs rather sparse, not in distinct rows. Scutellum with four mar- 
ginal bristles, anterior pair about half length of posterior ones. One 
humeral, two notopleurals. Mesopleura bare. No preapical tibial 
bristles. 

First vein ends near middle of wing. Auxiliary vein rudimentary, 
does not bend anteriorly at apex. Posterior crossvein present but 
very faint. Second vein strongly curved posteriorly in distal section. 
Discal and second basal cells separated. Anal crossvein faint. 
Costal index about 6, fourth vein index about 3. Costa reaches only 
to apex of third vein. 

Length, 2% mm., wing, 2mm. 

Type from Cottonwood Springs (near Mecca, Riverside County), 
Calif., May 1939 (USNM 61473). Sixteen paratypes from Cotton- 
wood Springs, Pasadena (M. R. Wheeler), and Lancaster, Calif., 
Prescott, Ariz. (M. R. Wheeler), and Mogollon, N. Mex. (M. R. 
Wheeler). Dr. Wheeler states that he also has collected this species 
at Peshashtin, Wash., Patagonia, Ariz., and Fort Davis, Tex. 


NEARCTIC PERISCELIDAE—STURTEVANT 555 


Exotic Periscelidae 


Two current keys to the genera of Periscelidae are confused; the key 
for the North American forms given by Curran (1934, p. 323), and the 
key for the genera of the world by Malloch (1932, p. 266). My own 
familiarity with exotic forms is limited to a single damaged specimen 
from Colombia in the collection of Cornell University. However, 
this specimen roused my suspicions of the validity of one of the major 
characters used in these keys. As a result, at my request C. W. 
Sabrosky has kindly examined the type specimens of Marbenia 
peculiaris Malloch, Neoscutops rotundipennis Malloch, Scutops fasci- 
pennis Coquillett, and S. maculipennis Malloch (all USNM) and re- 
ports that in all these the costa reaches only to the third vein. 

This is the case also with Periscelis (contrary to Curran’s key), 
which means that the first separation given by him is nonexistent if 
Cyamops is removed from the family. A further conclusion then be- 
comes evident: Panamenia Curran is a synonym of Scutops Coquillett, 
and P. chapman is doubtfully distinct from S. fascipennis Coquillett. 
The above-mentioned specimen from Colombia is also a Scutops, prob- 
ably best referred to S. fascipennis, though the wing pattern is slightly 
different. 

The account by Malloch (1932) is confused in that he says he had 
not seen the European Microperiscelis (Myodris), whereas the genotype 
is annulata, with which he was familiar. I think it was Periscelis that 
he had not seen. As indicated above, I do not think that Myodris is 
valid. The first separation in Malloch’s key is based on the anterior 
curvature of the auxiliary vein, and he places Scutops in the section 
without such curvature, which is contrary to the notes of Melander 
(1913a, p. 167) on the type specimen of S. fascipennis, the genotype. 
It may also be noted that utilization of this character makes Periscelis 
occidentalis, on Malloch’s key, run to the neighborhood of Marbenia, 
a genus that seems to me doubtfully distinct from Periscelis, though I 
am not prepared to insist on the point without seeing Marbenia. 

In view of the above it seems desirable to present a revised key to 
the genera of the world, even though I have seen only two of the five. 


Key to the genera of Periscelidae of the world 
(Modified from Malloch, 1932) 


Pee heirs Over entire surface! f! sow Slee eet. ee e--- 2 
Paes mare intcencral area. 2h VLU) 90) 10 OMT OD OOOO. 8. 3 
eves Of Conspicuous stalks_.__- 927°. ae Diopsosoma Malloch 
Pye not on stalks). p: oo. selene ete ee dees Neoscutops Malloch 

3. Face conspicuously flattened, more or less shield-shaped, and extending forward 
OSES CREO Pec) ee Oe a a ne eer Scutops Coquillett 


Face without a projecting shield-shaped area___-_-_-------------------- 4 


556 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


4. Face with two conspicuous transverse furrows-___-------- Marbenia Malloch 
Face not with two conspicuous transverse furrows__-------- Periscelis Loew 


List of the described Periscelidae of the world 


(Valid genera are in boldface. Synonyms are in italic. The first species listed 
under each genus is the genotype) 


Cyamops Melander, 1913b, p. 291. To Anthomyzidae. 
Diopsosoma Malloch, 1932, p. 267. 
D. prima Malloch, 1932, p. 267. Pert. 
Marbenia Malloch, 1931. p. 31. 
M. peculiaris Malloch, 1931, p. 32. Panamé. 
Meronychia Enderlein, 1917. Lapsus for Meronychina. 
Meronychina Enderlein, 1914, p. 327. Synonym of Periscelis. 
Microperiscelis Oldenberg, 1914, p. 39. Synonym of Periscelis. 
Myodris Lioy, 1864, p. 1103. Synonym of Periscelis. 
Neoscutops Malloch, 1926, p. 25. 
N. rotundipennis Malloch, 1926, p. 25. Costa Rica. 
Panamenia Curran, 1934, p. 323. Synonym of Scutops. 
Periscelis Loew, 1858, p. 118. Synonyms: Meronychina, Microperiscelis, Phorti- 
coides, Sphyroperiscelis, Myodris. 
P. annulipes Loew, 1858, p. 113. Europe. 
P. annulata (Fallén), 1813. (Notiphila.) Europe, United States. 
Myodris annulata Lioy, 1864, p. 1103. 
Microperiscelis annulata Oldenberg, 1914, p. 39. 
Meronychina annulata Enderlein, 1914, p. 327. 
Phorticoides flinti Malloch, 1915, p. 87. 
P. nebulosa Hendel, 1913, p. 389. Argentina. 
P. occidentalis, new species. Western United States. 
P. wheeleri (Sturtevant), 1923, p. 2. (Sphyroperiscelis.) United States. 
P. winnertzi Egger, 1862. Europe. 
Microperiscelis winnertzi Oldenberg, 1914, p. 39. 
Phorticoides Malloch, 1915, p. 87. Synonym of Periscelis. 
Podocera Czerny, 1929, p. 98. To Anthomyzidae. 
Scutops Coquillett, 1904, p. 97. Synonym: Panamenia. 
S. fascipennis Coquillett, 1904, p. 97. Central America. 
S. chapmani (Curran), 1934, p. 323 (=fascipennis Coquillett?). (Panamenia.) 
Panama. 
S. maculipennis Malloch, 1926, p. 24. México. 
Sphyroperiscelis Sturtevant, 1923, p. 1. Synonym of Periscelis. 


Family ANTHOMYZIDAE 


Since Cyamops Melander and Podocera Czerny, formerly referred 
to the Periscelidae, are being put in this family, an account of the 
Nearctic genera is presented. 

The following definition of the family is based largely on that of 
Collin (1944). 

Third antennal segment making nearly a right angle with the 
second; at least one reclinate orbital bristle; postverticals small and 
convergent, or absent; vibrissae present; no presutural dorsocentrals; 
mesopleura and disc of scutellum bare; first vein short, not over one- 
third length of wing; auxiliary faint apically, ending in costa just basal 


NEARCTIC PERISCELIDAE—STURTEVANT 557 


to first vein; distal costal break represented at least by a definite 
weakening; tibiae without preapicals. 

The antennal structure suggests that of the Periscelidae, but that 
family is easily distinguished from the Anthomyzidae by its long first 
vein; complete absence of distal costal break; divergent postverticals. 

The members of this family are sometimes treated as belonging to 
the Opomyzidae, which differ in having presutural dorsocentrals, 
hairy mesopleura, divergent postverticals, and no true vibrissae. 
The only Nearctic genus of Opomyzidae known to me is Geomyza, 
which has a large oral bristle that might be considered a vibrissa were 
it not situated some distance behind the anterior end of the row of 
oral hairs. The remaining genera listed by Curran (1934) under this 
family and not included in the key below make up the family Tethini- 
dae. (See Melander, 1951, p. 187.) 


Key to the Nearctic genera of Anthomyzidae 


1. Upper occiput convex; posterior margin of wing usually absent, making wing 


CLL sinh a1) pM pg ipl gc lg ihr Se a2 eo 1 Mutiloptera Coquillett 
Upper occiput concave; wing complete except sometimes with reduced anal 
EEC SSS Se SR A nc ves i ey ad SS Sy car ENR 2 

2. Postverticals small; 2 or 3 reclinate orbitals_—__.2././.-_.....---.------ 3 
Postverticals absent; usually only one conspicuous reclinate orbital; proclinate 
OLolvelsepresent OF sbsent. 4-20 .- ck s- ee eee eee Dee hate Se rage ere oem 4 

3. Second vein sinuate; marginal cell at level of posterior crossvein less than 
Bae win OF SUbMArEINAIL. - 20" (Fo e6 sn ketene ye Ischnomyia Loew 
Second vein greatly curved; marginal cell at level of posterior crossvein more 
pian ale width of submarginal —- 2222-22222 2.2 te Anthomyza Fallén 

4. Yellow species; anal angle of wing rudimentary ___-_-___ Stenomicra Coquillett 
avkiso ‘anal angle well:developed .. o 211-222 222. S222 S282 Yee. SIU 5 


5. Arista pubescent or short-plumose (branches not longer than length of third 
antennal segment); no comb of short spines on first femur; no proclinate 
SEND ee ee eee eens er er ee Mumetopia Melander 

Arista long plumose (branches longer than length of third antennal segment); 

a comb of short, close-set spines on distal inner flexor surface of first femur; 
aiprochnatelorbitallih. 2th 28) 220054 ab BOLa ie: Be 1 2 Cyamops Melander 

I have not seen Mutiloptera, which is perhaps an opomyzid. Of 
the other genera, the above key is based on examination of the indi- 
cated number of species (some Neotropical or Palearctic) of the follow- 
ing genera: [schnomyia, 2; Anthomyza, 6; Stenomicra, 1; Mumetopia, 

3; Cyamops, 2. 

Genus Cyamops Melander, 1913 


Brues and Melander (1932) and Curran (1934) included Cyamops 
in the Periscelidae. This genus was described as a “geomyzid,’’ and 
seems to me best placed in the Anthomyzidae, close to Mumetopia. 

In the keys in both the above works the Periscelidae are separated 
from the Anthomyzidae (treated as part of the Opomyzidae by 


558 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Curran) by the absence, in the former, of a break in the costa just 
before the apex of the first vein. In Cyamops there is no actual 
break here, but there is a definite weakening of the vein, differing, so 
far as I can see, in no way from that found in Mumetoyna. 

I have examined the internal female genitalia in Cyamops nebulosa 
Melander. There are three chitinized spherical spermathecae, of 
which two are attached to a common duct. The two spirally thick- 
ened ducts open into a large weakly chitinized pouch that arises from 
the dorsal surface of the uterus, thus somewhat suggesting the single 
common duct found in Periscelis. ‘There are two parovaria; no sperm 
or ventral receptacle was found. The other known Anthomyzidae 
(Anthomyza, Ischnomyia, and Mumetopia) differ in having only two 
spermathecae and no dorsal uterine pouch. The eggs of C. nebulosa 
are also unusual in that they have two short, thick, anterodorsal 
filaments suggestive of those found in Drosophila or Parallelomma. 
The eggs are white rather than dark blackish brown as in Periscelis. 

Since the genus may be looked for under the Periscelidae, and since 
I have available a new species and new data on the one previously 
described species, a synopsis is given here. 


Key to the species of Cyamops 


1. First tibiae and tarsi yellow, darkened only on terminal tarsal segment; second 
section of costa at least twice length of third section; two pairs of scutellar 


[Ou Nts 1) CGY erat a tM Ng ty li Me A ae etsy og C. nebulosa Melander 
First tibiae and tarsi largely black; second and third costal sections nearly 
equal in length; one pair of scutellar bristles__-__-_- C. imitata, new species 


Cyamops nebulosa Melander, 1913 


I have examined 43 specimens from Middleboro, Rochester, 
Mashpee, and Woods Hole, Mass., and Dismal Swamp (Cornell 
University collection) and Lake Drummond (H. S. Barber), Va. 
Specimens from Woods Hole (the type locality) are in the U. S. 
National Museum. The species is recorded by Johnson (1925) from 
Salisbury Cove, Maine, and Woburn, Mass. Mr. Sabrosky informs 
me that there are specimens in the U. 8S. National Museum from 
Beltsville and from near Lloyds (Dorchester County), Md., Alex- 
andria, Va. (bred May 20, 1951, from pupae collected by W. W. 
Wirth in osmundine bog), La Fayette, Ind., and Jacksonville, Fla. 

The original description was based on a male; the female differs in 
having a distinctly broader face. 

In eastern Massachusetts I have found the species easy to collect 
from June to the end of August by sweeping around the edges of 
shady sphagnum bogs. Of the 40 specimens so collected in this 
region, 16 (of both sexes) have no dark pattern on the wings. I had 
supposed these represented a distinct species, since their body color 


NEARCTIC PERISCELIDAE—STURTEVANT 559 


is not consistent with the hypothesis that they are teneral. However, 
through the kindness of Mr. Sabrosky I have seen a mated pair 
collected at Lake Drummond, Va. (H. S. Barber, June 8-11, 1905), 
in which the male has the typical pattern of wing clouds while the 
female has hyaline wings. It seems likely that we are concerned here 
with dimorphism of a single species, rather than with two species. 
Mr. Sabrosky reports that the other specimens listed above in the 
U.S. National Museum collection all have the typical wing pattern. 

C. nebulosa is variable in color. In some specimens the legs are 
wholly yellow, except that the terminal segments of all tarsi are 
darkened. In others there is more or less darkening of the apical 
portions of the second and third femora, which in some cases are as 
dark as in the single specimen of imitata. The leg color is not corre- 
lated with the clouding of the wings, and is less easily used for a sharp 
classification into distinct groups of individuals, since intermediates 
occur. 

Cyamops imitata, new species 


Female. Subshining black. Orbits silvery; antennae brownish. 
Legs, including coxae, yellow except brown to black first tibiae and 
tarsi, apical halves of second and third femora, and apices of second 
and third tarsi. Palpi and proboscis yellow. Knob of haltere white. 
Wing clouded, with a hyaline anterior border that includes costal and 
marginal cells and anterior half of submarginal, except that the apical 
portion of the latter is wholly clouded distal to a point midway between 
the ends of second and third veins. 

Arista with 8 or 9 long dorsal branches and 2 or 3 distal ventral 
ones; the 6 or 7 proximal dorsal branches are themselves bifid beyond 
their middle. One vibrissa; posterior to it are two downward-pointing, 
long bristles. Clypeus evident, small. A large proclinate orbital 
and a smaller reclinate one; no ocellars or postverticals; a single 
(divergent) vertical. Face narrowed by the eyes below the antennae. 
A single posterior dorsocentral; hairs on mesonotum sparse, not in 
regular rows; one supra-alar; one post-alar; no humerals; two noto- 
pleurals; mesopleura bare; one conspicuous sternopleural; a single 
(apical) pair of convergent scutellars; no preapicals on tibiae; a long 
apical on third tibia. Abdomen flattened. 

Second and third costal sections nearly equal; last section fourth 
vein about 2.3 times penultimate section. Third and fourth veins 
distinctly divergent apically. A conspicuous, erect bristle arising 
from costa near its base. 

Length, 2.5 mm. 

Type (USNM 61474) collected at La Fayette, Ind., July 6, 1915 
(J. M. Aldrich). 


560 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Genus Stenomicra Coquillett, 1900 


This genus was described as a drosophilid, and is still sometimes 
referred to that family. Sturtevant (1923) referred it to the ‘“‘Geo- 
myzidae,” and Hendel as well as Brues and Melander (1932) put it 
in the Asteidae, a reference not favored by Sabrosky (1943). Czerny 
(1929) described Podocera ramifera, from Ceylon, as a periscelid. 
Hendel and Malloch both point out that Podocera is a synonym of 
Stenomicra and that it should not be placed in the Periscelidae. 
Finally, the British and Fijian genus Diadelops Collin (1944, 1951) is 
evidently a synonym of Stenomicra. Collin places this genus in the 
Anthomyzidae, near Anagnota, a disposition that seems appropriate 
for Stenomicra. 

Stenomicra was overlooked by Curran (1934). Malloch (1927) 
has presented a key to the species of the world, of which Stenomicra 
angustata Coquillett is the only known American one. 


Stenomicra angustata Coquillett, 1900 


Coquillett (1900) recorded this species from Puerto Rico, and 
Sturtevant (1923) added localities in Pennsylvania, Florida, Alabama, 
and Louisiana. I have since seen four specimens collected by Dr. H. D. 
Stalker at Creve Coeur, Mo., about the reedy shores of a fresh-water 
pond (July 15, 1951). 


References 


Bruss, C. T., and MELANDER, A. L. 
1932. Classification of insects. Bull. Mus. Comp. Zool., vol. 73, 672 pp. 
Coun, J. E. 
1944. The British species of Anthomyzidae (Diptera). Ent. Monthly 
Mag., vol. 80, pp. 265-272. 
1951. A new species of Diadelops Collin (Diptera: Anthomyzidae) from 
Fiji. Proc. Roy. Ent. Soc. London, Ser. B, vol. 20, pts. 3-4, pp. 
47-48. 
CoquiL.LeTt, D. W. 
1900. Report on a collection of dipterous insects from Puerto Rico. Proce. 
U.S. Nat. Mus., vol. 22, pp. 249-270. 
1904. New Diptera from Central America. Proc. Ent. Soc. Washington, 
vol. 6, No. 2, pp. 90-98. 
Curran, C. H. 
1934. The families and genera of North American Diptera, 512 pp. 
Czrerny, A. L. 
1929. Podocera ramifera, eine neue Gattung und Art der Perisceliden von 
Ceylon. Konowia, vol. 8, pt. 1, pp. 93-94. 
EaGGErR, JOHANN 
1862. Dipterologische Beitrage. Verh. Zool.-Bot. Ges. Wien, vol. 12, pt. 
3, pp. 777-784. 


NEARCTIC PERISCELIDAE—STURTEVANT 561 


ENDERLEIN, GUNTHER 
1914. Diptera, Fliegen, Zweifliigler, in Brohmer, Fauna von Deutschland, 
pp. 313-368. 
1917. Dipterologische Studien. XVI. Dipterologische Notizen. Zool. 
Anz., vol. 49, Nos. 3-4, pp. 65-72. 
Fautén, Cart F. 
1813. Berskrifning 6fver nagra i sverige fume vattenflugor. Svenska Ve- 
tensk. Akad. Handl., vol. 34, pp. 240-257. 
HENDEL, F. 
1913. Neue Drosophiliden aus Siidamerika und Neuguinea (Dipt.). Ent. 
Mitt., vol. 2, pp. 386-390. 
JOHNSON, CHARLES W. 
1925. Fauna of New England. 15. List of the Diptera or two-winged 
flies. Occas. Papers Boston Soc. Nat. Hist., vol. 7, 326 pp. 
Lioy, PaoLo 


1864. I Ditteri . . . , Atti Ist. Venuto, vol. 9, ser. 3, pp. 1087-1126. 
Lorw, H. 
1858. Ueber einige neue Fliegengattungen. Berliner Ent. Zeitschr., vol. 2, 
pp. 101-122. 
Mattocg, J. R. 


1915. An undescribed sapromyzid (Diptera). Bull. Brooklyn Ent. Soc., 
vol. 10, No. 4, pp. 86-88. 
1926. New genera and species of acalyptrate flies in the United States 
National Museum. Proc. U.S. Nat. Mus., vol. 68, art. 21, 30 pp. 
1927. The species of the genus Stenomicra Coquillet (Diptera, Acalyptrata). 
Ann. Mag. Nat. Hist., ser. 9, vol. 20, pp. 23-26. 
1931. Notes on some acalyptrate flies in the United States National Museum 
Proc. U. S. Nat. Mus., vol. 78, art. 15, 32 pp. 
1932. A new genus of diopsid-like Diptera (Periscelidae). Stylops, vol. 1, 
pt. 12, pp. 266-268. 
MELANDER, A. L. 
1913a. Some acalyptrate Muscidae. Psyche, vol. 20, pp. 166-169. 
1913b. A synopsis of the dipterous groups Agromyzinae, Milichiinae, Ochthi- 
philinae and Geomyzinae. Journ. New York Ent. Soc., vol. 21, 
No. 4, pp. 283-300. 
1951. The North American species of Tethinidae (Diptera). Journ. New 
York Ent. Soc., vol. 59, No. 4, pp. 187-212. 
OLDENBERG, LORENZ 
1914. Beitrag zur Kenntnis der europiischen Drosophiliden (Dipt.). 
Arch. Naturg., vol. 80A, pt. 2, pp. 1-42. 
Sasprosky, C. W. 
1943. New genera and species of Asteiidae (Diptera), with a review of the 
family in the Americas. Ann. Ent. Soc. Amer., vol. 36, No. 3, pp. 
501-514. 
Sturtevant, A. H. 
1923. New species and notes on synonymy and distribution of Muscidae 
Acalypteratae. Amer. Mus. Nov., vol. 76, 12 pp. 
1926. The seminal receptacles and accessory glands of the Diptera with 
special reference to the Acalypterae. Journ. New York Ent. Soc., 
vol. 34, No. 1, pp. 1-21. 


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PROCEEDINGS OF THE UNITED STATES .NATIONAL MUSEUM 


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SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1954 No. 3333 


NORTH AMERICAN TRICLAD TURBELLARIA, XIII: 
THREE NEW CAVE PLANARIANS 


By Lissir H. Hyman 


Previously (Hyman, 1937, 1939, 1945) I have described a total of 
11 species of planarians from the waters of North American caves. 
Comment was made in these articles on the extensive speciation ex- 
hibited by the cave planarians of the United States. It was pointed 
out that every locality appears to have a different species of cave 
planarian and that the investigation of caves in new localities in- 
evitably reveals new species of these animals. The present report is 
based on collections from caves in three new localities and each lo- 
eality is again found to harbor a new and different species of cave 
planarian. As generic and familial definitions have been presented in 
my previous articles on cave planarians, it appears unnecessary to 
repeat them here. 


Family PLANARIIDAE 
Phagocata cavernicola, new species 


Figure 65,a-c 


Material.—Five specimens sent by the U. S. National Museum 
(USNM). 

Form.—Small, slender, elongate (fig. 65, a), around 7 to 8 mm. long, 
probably up to 10 mm. in length; anterior end appears slightly rounded 
as in figure 65,a in some of the specimens, truncate or slightly in- 
dented in others (fig. 65, 6). 

Color.—White in life, according to the collector, but turns brown on 

preservation. 

) 290841—54 563 


564. PROCEEDINGS OF THE NATIONAL MUSEUM vot. 103 


Eyes.—Typically two, but two of the five specimens show super- 
numerary eyes as in figure 65, a, 6; possibly this indicates a breakdown 
of the eyes in relation tothe cave habitat. 

General structure—The histological condition of the specimens is 
poor and it is impossible to furnish details of the structure. The 
pharynx appears somewhat long for the size of the animal (fig. 65, a). 
In the median line of the head anterior to the eyes a light streak was 
noticeable on the whole animals (fig. 65, a) but one of the heads was 
sectioned sagittally and, although its histological condition is bad, it 
appears certain that the midregion does not differ histologically from 
the rest of the head. The body margins do not present any enlarged 
rhabdites. 

Reproductive system.—Testes evident in the worms when whole and 
extend in lateral regions from the level of about the fourth intestinal 
diverticulum to the level of the gonopore (fig. 65, a). Transverse 
and sagittal sections show that the testes are situated ventrally. 
A sagittal view of the copulatory apparatus, constructed from two 
sets of sagittal sections, is shown in figure 65, c. The sperm ducts, 
ascending from below into the slightly developed penis bulb, enter 
separately the bulbar cavity. This is an oval cavity that makes a 
right-angled bend and continues through the penis papilla as the 
ejaculatory duct. The penis papilla has a short truncate form, and is 
noticeably lacking in muscularity. It is also asymmetrical, very 
decidedly so in one of the sagittal series; its dorsal wall is longer than 
the ventral wall. The penis papilla lies in a well-developed male 
antrum that narrows towards the common gonopore. Juxst before 
joining the vagina, the male antrum receives the common ovovitelline 
duct into its dorsal wall. The female canal shows an unusual degree 
of separation from the male antrum. ‘There is, in fact, no develop- 
ment of a female antrum or common antrum as the vagina joins the 
male antrum almost at the gonopore. The vagina or terminal part of 
the female canal is somewhat expanded and lined by a very tall 
epithelium. The vagina is continuous with the bursal canal of which 
it is really the terminal portion. The bursal canal as usual is a narrow 


ExpLaNaTION OF FIGURES 


1, eyes 15, cavity of adhesive organ 

2, intestinal diverticula 16, eosinophilous lining of adhesive organ 
3, testes 17, retractor muscle of adhesive organ 
4, pharynx 18, adhesive organ 

5, sperm ducts 19, ovaries 

6, penis bulb 20, margin of large rhabdites 

7, penis papilla 21, adhesive tail region 

8, ejaculatory duct 22, eosinophilous gland cells 

9, male antrum 23, protractor muscle 

10, vagina 24, gland areas of male apparatus 

11, copulatory bursa 25, female antrum 

12, bursal canal 26, common antrum 


13, entrance of ovovitelline duct 27, common gonopore 
14, bulbar cavity : 


THREE NEW CAVE PLANARIANS—HYMAN 565 


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Ficure 65.—a, General view of Phagocata cavernicola; b, anterior end of another specimen 
of P. cavernicola, showing supernumerary eyes; c, sagittal view of the copulatory apparatus 
of P. cavernicola, posterior end above; d, sagittal section of the adhesive organ of Sphal- 
loplana georgiana. (For explanation see facing page.) 


566 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


duct passing anteriorly above the male apparatus and terminating in 
a rather small sac, the copulatory bursa, situated immediately anterior 
to the penis bulb, between this and the posterior end of the pharyn- 
geal chamber (fig. 65, c). The entire female copulatory apparatus 
(copulatory bursa, bursal canal, and vagina) is but slightly provided 
with musculature. 

Differential diagnosis—-This species differs from other North 
American species of Phagocata in the asymmetrical penis papilla, 
expansion of the terminal part of the bursal canal into a vagina, lack 
of a female antrum, and sharp separation of the vagina from the male 
antrum. 

Locality.—Collected by R. C. Hoffmaster in 1949 and again on 
Jan. 19, 1951, in Evac Cave near Hillside, Pa. An additional small 
specimen from Conodoguinet Cave in the same region, also collected 
in 1949, is presumably the same species although this could not be 
determined with certainty, as the specimen is immature. 

Holotype——One whole mount deposited in the U. S. National 
Museum (USNM 24610); also set of transverse sections (4 slides) 
and set of sagittal sections (8 slides) in this institution. 

Remarks.—The occurrence of eyes in a cave planarian is always 
unexpected but eyes are present in two other North American cave 
planarians, namely, Phagocata subterranea Hyman, 1937, and Soro- 
celis americana Hyman, 1939b. Thus both of our cave planarians 
that belong to the genus Phagocata have retained their eyes and this 
may indicate only a moderate degree of adaptation of this genus to 
the cave habitat. It is to be noted, further, that there are white 
epigean species of Phagocata in the United States, so the absence 
of pigment in the cave species of this genus cannot be regarded as 
adaptive. Sorocelis americana often occurs in epigean habitats, so 
its retention of eyes is not surprising. 


Family KENKIIDAE 
Sphalloplana georgiana, new species 
Fiauress 65,d; 66,6; 67,a 


Material.—Four specimens presented by C. E. Mohr. 

Form.—Elongated, slender but less so than the preceding species; 
narrowed anteriorly with truncate anterior margin bearing a central 
adhesive organ (fig. 66, 6); posterior end bluntly pointed; about 8 mm. 
long. 

Color.—White. 

Eyes.—W anting. 

General structure—The histological condition of the specimens, 
three of which were sectioned, is very bad. The pharynx appears 
relatively small compared to its size in the preceding species. The 
digestive diverticula are exceedingly numerous and narrow as indi- 


Ficure 66.—a, 


ae 


VETS 


General view of Speophila hoffmasteri; b, general view of Sphalloplana 
georgiana. (For explanation see page 564.) 


568 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


cated in figure 66,b. In the case of white planarians with truncate 
anterior margin, it is necessary to examine transverse sections of the 
pharynx to determine the family. Such sections were prepared but 
because of their bad histological condition I was left in some uncer- 
tainty on this point. As far as I can make out, however, the arrange- 
ment of the inner pharynx musculature corresponds to the family 
Kenkiidae but the possibility remains that the animal might belong 
to the Dendrocoelidae, which it much resembles externally. The 
body margins are provided with enlarged rhabdites as characteristic 
of cave planarians. 

Adheswe organ.—An adhesive organ is found in the center of the 
anterior margin in all members of the family Kenkiidae and also 
frequently in members of the Dendrocoelidae. Sagittal sections of 
the anterior end of Sphalloplana georgiana showed a rather simply 
corstructed adhesive organ as characteristic of the genus. This 
region of the specimens proved to be in better histological condition 
than the rest of the body. A sagittal view of the adhesive organ is 
given in figure 65,d. There is a small cuplike depression at the 
margins of which the regular covering epidermis ceases rather abruptly, 
being replaced by a noncellular margin dotted with eosinophilous 
secretion. From the inner surface of the cup longitudinal retractor 
muscles proceed posteriorly and curve ventrally to join the regular 
ventral subepidermal musculature. The eosinophilous glands, which 
presumably provide the eosinophilous secretion, could not be found 
in the available material. 

Reproductive system.—The testes could not be seen in any of the 
animals when whole but were discovered in sections in a poor condi- 
tion in two rather short, lateral bands immediately anterior to the 
pharynx and have been inserted in figure 66,6. As far as could be 
ascertained, they do not extend anteriorly as far as is usually the 
case in planarians. The testes are small and situated ventrally. The 
ovaries were not discernible in the sections. The copulatory appa- 
ratus is situated well behind the pharynx as shown in figure 66,0. 
It was in bad condition in both of two sets of sagittal sections pre- 
pared, but a sagittal view, given in figure 67,a, has been pieced to- 
gether by study of these sections. There is a well-developed, rounded, 
and highly muscular penis bulb that was conspicuous in the whole 
animals (fig. 66, 6) but less evident in sections. The sperm ducts, 
ascending from below, pass separately through the penis bulb and 
open into the sides of the rounded bulbar cavity, occupying the cen- 
ter of the bulb (fig. 67, a). The bulbar cavity continues as a rather 
wide ejaculatory duct through the conical penis papilla to the tip of 
the latter. The penis papilla is somewhat muscular and clothed with 
a tall epithelium. It lies in a spacious male antrum, into the dorsal 
wall of which there opens the common ovovitelline duct surrounded 


569 


THREE NEW CAVE PLANARIANS—-HYMAN 


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Ay 


AZOAN 
Ti siiwy SOS 
Wf PYMENWATAY 


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Sagittal view of the copulatory apparatus of Sphalloplana georgiana, anterior 


—4, 


Ficure 67 


sagittal section of the adhesive organ of Speophila hoffmastert. (For explana- 


tion see page 564.) 


b, 


° 
? 


end above 


570 PROCEEDINGS: ‘OF. THE NATIONAL MUSEUM VOL. 108 


by eosimophilous glands. The copulatory bursa could not be found 
on any of the available sections but its slender canal was more or 
less in evidence, passing dorsal to the male apparatus. This curves 
downward and enters the dorsal posterior angle of the greatly ex- 
panded female antrum, which exits below by the common gonopore 
and receives the male antrum into its anterior wall. The walls of 
the female antrum are greatly thickened, especially dorsally. This 
thickening consists mainly of muscle fibers, chiefly circular, but the 
thick dorsal wall seems to contain, next to the lining epithelium, a 
glandular layer through which ual fibers pass. 

Differential diagnosis.—Sphalloplana georgiana differs from other 
species of the genus in the greatly expanded female antrum with 
thick walls, presumably serving as a vagina. 

Locality.—Collected by C. E. Mohr in Waterfall Cave, Trenton, 
Ga., Dec. 8, 1950. 

Holotype.—The best of the specimens has been mounted whole as 
a holotype and deposited in the U. S. National Museum (USNM 
24614). However, as the anterior margin of this specimen appears 
damaged, a whole mount of the anterior part of another specimen 
showing the adhesive organ is also deposited. 


Speophila hoffmasteri, new species 
FIGURES 66,a; 67,b; 68 


Material—Two specimens sent by the U. S. National Museum 
and one specimen presented by Leslie Hubricht. 

Form.—Size moderate, around 11 mm. in length, plump, narrowed 
at the ends; anterior margin rounded with an adhesive organ (fig. 
66, a). 

Color.—White. 

Eyes.—Wanting. 

General structure.—The histological condition of the worms is fair. 
The pharynx appears very short for the length of the specimens (fig. 
66,a) but may be contracted. The intestinal diverticula were not 
clearly evident on the whole mount. The body margin is provided 
with a thick zone of enlarged rhabdites and this widens considerably 
on the tail end, which therefore must be highly adhesive. Such a 
wide marginal zone of large rhabdites is characteristic of cave plana- 
rians of the family Kenkiidae and gives these worms unusual ability 
to cling to‘ objects. 

Adhesive organ.—At the anterior margin there is a conspicuous and 
well-developed adhesive organ of the type characteristic of the genus. 
It is shown in surface view in figure 66,a, and in sagittal section in 
figure 67,6. The surface epithelium stops as usual at the lips of the 
organ which is lined throughout with an indefinite layer filled with 


571 


THREE NEW CAVE PLANARIANS-~HYMAN 


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Ficure 68.—Sagittal view of the copulatory apparatus of Speophila hoff masieri, anterior 


(For explanation see page 564.) 


end above. 


572 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


eosinophilous granulations. The organ has the form of a long tubular 
gland whose lumen is filled with eosinophilous secretion. At about 
the middle of the gland are found lateral outpocketings, and from 
this point posteriorly there are seen to either side of the gland the 
long-necked eosinophilous glands that furnish the secretion. They 
appear as darkly stained pyriform bodies. At the posterior end of the 
gland a retractor muscle extends posteriorly, joining the muscle 
layer of the ventral body wall, and a thicker and more prominent 
protractor muscle curves dorsally and anteriorly to insert on the dorsal 
body wall. The presence of a protractor muscle indicates that the 
gland can be everted to some extent, probably as far as the lateral 
pouches. The eosinophilous secretion is of an adhesive nature and 
it is generally supposed that the adhesive organ functions in the 
capture of prey as well as in leechlike crawling. 

Reproductive system.—The gonads could not be seen in whole speci- 
mens but were found in the sections and have been entered on figure 
66,a. The pair of ovaries occurs in the usual site. The testes form 
a tract on either side anterior to the pharynx. They are rather large 
and fill the middle region of the sections. The copulatory apparatus 
was in poor condition in both sets of sagittal sections that were pre- 
pared. The copulatory bursa was made out with great difficulty 
and the dorsal wall above the vagina was badly broken in both series. 
A sagittal view of the copulatory apparatus as constructed by study 
of the two series of sections is given in figure 68. The penis is a con- 
spicuous object behind the pharynx in the whole worm (fig. 66,a) 
but as the penis bulb appears somewhat curved its full extent is not 
evident in any one section. There seems to be an unusually large and 
muscular penis bulb formed of muscle fibers paralleling its contours 
and penetrated by the two narrow sperm ducts (fig. 68). At the base 
of the penis papilla these join to a narrow ejaculatory duct that soon 
widens as it traverses the penis papilla, narrowing again towards the 
tip of the latter. The penis papilla is of elongated conical form and 
rather muscular. It presents the peculiarity of a layer of cyanophilous 
gland cells outside the lining epithelium of the ejaculatory duct along 
the distal part of the latter. The penis bulb lies within a male antrum 
that follows its contours. As already indicated, the female apparatus 
was in poor condition in the available material. The copulatory 
bursa was much damaged but appeared as indicated in figure 68 as a 
flattened sac with a low epithelial lining in its anterior half, a tall 
lining posteriorly where the bursa passes into its canal. The latter 
is a long, slender duct proceeding posteriorly above the male antrum. 
After reaching a point behind the level of the gonopore, the bursal 
canal turns abruptly ventrally and enters a conspicuous, rounded 
vagina that was very evident in the whole specimens (fig. 66,q). 


THREE NEW CAVE PLANARIANS—HYMAN 573 


The vagina is lined by a very tall epithelium and this is surrounded 
by a wide halo of what seems to be mesenchyme traversed by radial 
muscles. Distally the vagina narrows to a short tube that joins the 
male antrum, forming a very small common antrum that opens below 
by the common gonopore. The entrance of the ovovitelline duct 
into the copulatory apparatus could not be found but presumably 
this duct opens into the roof of the male antrum. In the posterior 
wall of the male antrum is found an area of cyanophilous glands 
similar to those surrounding the ejaculatory duct in the penis papilla. 

Differential diagnosis.—Speophila hoffmasteri is distinguished from 
the other known species of the gerus by the round, thick-walled 
vagina and the layer of gland tissue in the penis papilla and wall of 
the male antrum. 

Locality.—Collected by R. E. Hoffmaster in Blowing Cave, Pendle- 
ton County, W. Va. (no date), and by Leslie Hubricht in Mystic Cave, 
Pendleton County, W. Va., May 30, 1952. 

Holotype-—One whole mount of the better of the Hoffmaster 
specimens deposited in the U. S. National Museum (USNM 24616); 
also one set of sagittal sections to show the adhesive organ (5 slides) 
and one set of sagittal sections of the copulatory region (3 slides) 
in the same institution. 


References 
Hyman, LissprE HENRIETTA 

1937. Studies on the morphology, taxonomy, and distribution of North 
American triclad Turbellaria. VIII. Some cave planarians of the 
United States. Trans. Amer. Micr. Soc., vol. 56, pp. 457-477, 
21 figs. 

1939a. New species of flatworms from North, Central, and South America. 
Proc. U. S. Nat. Mus., vol. 86, pp. 419-439, 5 figs. 

1939b. North American triclad Turbellaria. X. Additional species of cave 
planarians. Trans. Amer. Micr. Soc., vol. 58, pp. 276-284, 9 figs. 

1945. North American triclad Turbellaria. XI. New, chiefly cavernicolous, 
planarians. Amer. Midl. Nat., vol. 34, pp. 475-484, 8 figs. 


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niftqaqseisnd skh oh toub raoisionata adi yaibapnosy 
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1954 No. 3334 


A SUPPLEMENT TO W. M. TATTERSALL’S REVIEW OF THE 
MYSIDACEA OF THE UNITED STATES NATIONAL MUSEUM! 


By AvsBert H. BANNER” 


In 1942 Dr. Walter M. Tattersall completed the manuscript of his 
lengthy study of the crustaceans of the order Mysidacea of the U.S. 
National Museum. This study, more than a simple catalog of speci- 
mens stored at the museum, was a listing of all species of the order 
ever reported from the Americas, both North and South, and a review 
of all literature pertinent to them. Many species were described and 
depicted, taxonomic confusions were cleared, and 1 new genus and 19 
new species were described. 

This study, as it stands, is the most valuable contribution to the 
knowledge of the order yet made in America and will be invaluable to 
every worker, specialist or not, who will need to identify these crus- 
taceans. However, Dr. Tattersall died on October 1, 1943, before 
even the initial preparations were made by the U.S. National Museum 
for the publication of the study. Because of the war-induced delay, 
the manuscript was not finally published until October 4, 1951. In 
the intervening eight years several papers on the group appeared, and 
others, published earlier in Europe during the war, became available 
for the first time. Some of these contain references to the species 
discussed by Tattersall, and a few changes in taxonomic status have 
been introduced. As references to these additional studies were not 
inserted in the manuscript in the course of editing, the actual date of 
publication is deceptive. 


1Tattersall, Walter M., A Review of the Mysidacea of the United States National Museum. U. 5. 
Nat. Mus. Bull. 201, x + 292 pp., 103 figs., Oct. 4, 1951. 
2 Department of Zoology and Entomology, University of Hawaii, Honolulu, T. H. 


290840—54- 575 


576 | |. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


As author of one of the pertinent papers mentioned above, I was 
asked by Dr. Fenner A. Chace, Jr., of the U. S. National Museum, 
and by Mrs. Olive S. Tattersall to prepare this short paper to supply 
the information needed to bring Dr. Tattersall’s study up to its actual 
date of publication. I wish to thank Dr. Chace and Mrs. Tattersall 
for asking me to make this emendation. 

In preparing these addenda, I hope.also to keep the seeming errors, 
resulting from the long delay in printing, from reflecting in any way 
on the excellent reputation of Dr. Tattersall in the eyes of those who 
may not know the true cause of the omissions, and to keep these 
omissions from confusing later workers, especially those nonspecialists 
who may not be in a position to investigate all the literature on the 
group. : | . 

No more will be done than to point. out under the various species 
the additions to the literature pertinent to their taxonomic status or 
to the American records. Where there may be a question about the 
taxonomy of any species, it will merely be mentioned but not resolved; 
finally, any publications issued after the date of publication of Tatter- 
‘sall’s work will not be reported. 


7 Addenda 
Page 3. 
The list of American species should be changed as follows: 
Baplamily Boreomysinae.. 2-2 +5 - scone a ene = 8 species 
‘rine ry chropuitr. 22 Sey 7 iy | eee ee 28 
Tribe Mysini..-.....5-.-2.4<:-..... Crm Aa 35 
Subfamily! Mysidellinaes !2v- s2ccn- ol} 2t- 2boests-2iee--eRee 1 
Page 4. 


To the fresh-water species of mysids should be added Neomysis mer- 
cedis, from the west coast of North America (Banner, 1948, pt. IT, 
Dp. (3). 

The total of littoral species for the Pacific coast should be raised to 
32 and Mysis oculata should be added as a species common to both 
coasts of North America (see the discussion under that species below). 


Page 5. 
In table 1 the number of west coast species of Acanthomysis should 
be changed to 9. 


Page 7. 

To the list of deeper-water species found only off the continental 
shelf of the Pacific slope should be added Katerythrops sp. and Mysidella 
americana. tf sien ; 

To the list of pelagic species common to both the Atlantic and Pacific 
coasts should be added Boreomysis microps. 


SUPPLEMENT TO TATTERSALL’S MYSIDACEA—BANNER 577 


Page 8. 

To the list of pelagic species known only from the Pacific side should 
be added Boreomysis kincaidi, Euchaetomeropsis pacifica, and Caesaro- 
mysis vanclevei, while Caesaromysis sp. should be substituted for 
Caesaromysides liguriae. 

In table 2, Neomysis mercedis should be listed as a species distrib- 
uted from Alaska to California rather than from British Columbia to 
California. (The identification of the Alaskan specimens was some- 
what uncertain.) 

Page 14.—Chalaraspidum alatum (Willemoes-Suhm) 

Fage (1941, p. 4) not only discussed the anatomy and relationships 
of this species but also recorded its capture from the Pacific Ocean 
near Panama. 


Pages 15-22.—Lophogaster M. Sars 


Any worker dealing with the genus Lophogaster should also consult 
the work of Fage (1942, pp. 1-39) in which, like Tattersall, he re- 
viewed the known species and described new species. Fage, more- 
over, established new criteria, not used by Tattersall and previous 
workers, which serve to distinguish the species. 

Below is a listing of the species of Lophogaster recognized by Tatter- 
sall and Fage, those arranged in opposition being presumably the same. 
In Fage’s list, the species prefaced by an asterisk are those for which 
he gave additional records, some of which are American. 


TATTERSALL FaGE 
L. rotundatus Nig *I. rotundatus 
*L. typicus 
L. typicus M. Sars *I. subglaber Hansen 3 
*L. challengeri Fage 
L. americanus Tattersall 
L. spinosus Ortmann *L. spinosus 
L. hawaiensis Fage *[. hawaiensis 
L. longirostris Faxon L. longirostris 
*L. pacificus Fage 
L. intermedius Hansen L. intermedius 


*L. multispinosus Fage 
*L. schmitti Fage 
L. japonicus Tattersall 
L. erythraeus Colosi L. typica var. erythraeus 
L. affinis Colosi L. subglaber var. affinis 
On the basis of the published descriptions it would appear that the 
new species described by Tattersall (LZ. americanus and L. japonicus) 
are not synonyms of the six described or reestablished by Fage (L. sub- 
glaber, L. challengeri, L. hawaiensis, L. pacificus, L. multispinosus, 


3 Nouvel (1943, p. 90) described L. subglaber insulare a new variety from the eastern Atlantic (referred to 
earlier in his paper as L. challengeri Fage). 


‘578 =~ /PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


and L. schmitt). However, final judgment on the validity of Tatter- 
sall’s species will have to wait until the specimens are examined. 


Pages 25-32.—Gnathophausia Willemoes-Suhm 

Like the species of the genus Lophogaster, the species of this genus 
were reported on by Fage (1941) on the basis of the extensive Dana 
collections. In his work Fage discusses both the morphology of the 
species and their distributional patterns. 


Page 25.—Gnathophausia ingens (Dohrn) 

This species was discussed by Fage (1941, p. 15) and many new 
records were given for it, including some from the Caribbean and 
adjacent waters. 


Page 26.—Gnathophausia gigas Willemoes-Suhm 

Records of this species, including some from off Central America, 
were given by Fage (1941, p. 24). Nouvel (1943, p. 12) gave some 
records from off Newfoundland. In addition, new records from off 
the northwestern coast of North America were given in my paper 
(Banner, 1948, pt. II, p. 358). 


Page 28.—Gnathophausia gracilis Willemoes-Suhm 

Records of this species from off Central America were given by 
Fage (1941, p. 27). 
Page 29.—Gnathophausia zoea Willemoes-Suhm 

One record was given by Fage (1941, p. 34) from off the northeastern 


coast of South America; Nouvel (1943, p. 15) gave two records of 
capture from off Newfoundland. 


Pages 32-34.—Eucopia Dana 

Three papers dealing with the names applied to the species and the 
separation of the species of this genus were published while Tattersall 
was finishing his manuscript. The studies were those of Nouvel 
(1942a, 1943) and Fage (1942). 
Page 32.—Eucopia sculpticauda Faxon 

Fage (1942, p. 56) recorded many captures of this species, including 
one off the eastern coast of the United States, two in the Caribbean 
Sea, and several off the Pacific coast of Central America and northern 
South America. 


-Page 33.—Eucopia australis Dana 

' Fage (1942, p. 41) recorded additional distributional records for 
this species, including two captures on the Pacific side of Panama and 
one in the eastern Caribbean. 


SUPPLEMENT TO TATTERSALL’S MYSIDACEA—BANNER 579 


Page 33.—Eucopia major Hansen 

Fage (1942, p. 41 ff.) has placed this species in the synonymy of LF. 
australis; Nouvel (1943, pp. 95-97) rejected the proposal and retained 
E. major as a separate and distinct species. Nouvel. (p. 28) 
gave one record from off Newfoundland in addition to non-American 
collections. 

Page 34.—Eucopia unguiculata (Willemoes-Suhm) 

This species was divided into E. hanseni and E. grimaldii, two new 
species, by Nouvel (1942a, pp. 3-6). This distinction was accepted 
by Fage (1942, p. 40) but he used the names only provisionally until 
an examination of the type should determine which form must retain 
the name FE. unguiculata. Tattersall and Tattersall (1951, p. 99) 
established that the name EH. unguiculata should be retained for 
E. hanseni Nouvel and that FE. grimaldii Nouvel is to be accepted. 
Unless subsequent work makes it necessary to reject Nouvel’s separa- 
tion, the specimens recorded by Tattersall should be reexamined and 
separated into the new species.‘ 

Additional records for the two species were given by Fage (1942, 
p. 47), including some records from near the coasts of the Americas, 
by Nouvel (1943, pp. 30-43), where there is one record from off New- 
foundland, and by Banner (1948, pt. I, p. 359), including many 
records from off the northwestern coast of North America. 


Page 45.—Boreomysis G. O. Sars 

Instead of about 20 species belonging to this genus, as given by 
Tattersall, there are now about 25 or 26 species recognized; only one 
of these new species has been described from American waters. 
Page 48.—Boreomysis tridens G. O. Sars 

Nouvel (1942b, p. 1; 1943, p. 45) described a variety lobata as new 
from off Newfoundland. 
Page 49.—Boreomysis arctica (Krgyer) 

Add synonym: B. tregonboffi Bacesco (1941a, p. 168; 1941b, p. 12) 
[according to Nouvel, 1943, p. 52]. 
Page 52.—Boreomysis californica Ortmann 
_A description, figures, and additional records from off the north- 
western coast of North America were given by Banner (1948, pt. I, 
p. 367). 
Page 55.—Boreomysis microps G. O. Sars 

Additional records for this species were given by Nouvel (1943, 
p. 48), including one record from off Newfoundland, and by Banner 
(1948, pt. I, p. 365) from off the northwestern coast of North America. 


41 plan to publish in the near future a review of the names and the specific distinctions applied within 
this genus. 


580 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Page 60. 


To the American species of Boreomysis should be added the species 
B. kineaidi Banner (1948, pt. I, p. 362) with a range from Washington 
to Alaska.5 


Page 60.—Siriella thompsonii (H. Milne-Edwards) 


Perhaps reference here should be made to the records of Nouvel 
(1943, p. 62), some of which are from locations that lie between the 
Azores and Newfoundland. 


Page 81.—Archaeomysis grebnitzkii (Czerniavsky) 


This species also has been described and depicted by Banner 
(1948, pt. I, p. 369, pl. 5). The records are for the ocean coast and 
the inland waters of Washington, the middle of the range established 
by Tattersall. 


Page 100.—Anchialina typica (Kr¢yer) 


This species was recorded from American waters south of New- 
foundland by Nouvel (1943, p. 70). 


Page 106.—Holmesiella anomala Ortmann 


This species has been redescribed and discussed with additional 
records (Banner, 1948, pt. I, p. 395). It should be noted that both 
Tattersall and Banner observed that there are two races of the species 
that reach different sizes at maturity. 


Page 112.—Euchaetomera tenuis G. O. Sars 


To the records of this species should be added those collections 
made off British Columbia (Banner, 1948, pt. I, p. 383), a considerable 
extension of the range as given by Tattersall. 


Page 112. 


After the genus Huchaetomera should be added the genus and 
species Huchaetomeropsis pacifica Banner (1948, pt. I, p. 285), a new 
species described from British Columbia and the first record of the 
genus from American waters of either coast. 


Page 113.—Meterythrops robusta S. I. Smith 

Additional records for this species have been given from Wash- 
ington to Alaska (Banner, 1948, pt. I, p. 377). This paper also noted 
that there were small-eyed individuals similar to those described by 
Tattersall as the new species M. microphthalma, but that after the 
examination of over 85 specimens the conclusion was reached that 
differences between the two forms were bridged by intermediate 
forms. If M. microphthalma is considered to be a synonym of M. 


5 I plan to discuss B. californica and B. kincaidi in a subsequent paper. 


SUPPLEMENT TO TATTERSALL’S MYSIDACEA—BANNER 58] 


robusta, then the record given for the new species constitutes the 
furthest extension of the range for M. robusta. 


Page 118.—Katerythrops Holt and Tattersall 


To the species of the genus should be added Katerythrops sp. 
Banner (1948, pt. I, p. 379), which is the only record of the genus in 
the Pacific. 


Page 120.—Longithorax capensis Zimmer 


Tattersall states that his specimens from Bermuda are the only ones 
known save the type; however, Nouvel (1943, p. 75) has recorded 
from the Azores a damaged specimen probably of this species. 


Page 128.—Amblyops abbreviata (G. O. Sars) 
To Tattersall’s record of this species from Alaska should be added 


the two records from Alaska and one record from British Columbia 
(Banner, 1948, pt. I, p. 382). 


Page 134.—Pseudomma truncatum S. I. Smith 


To the west coast records of this species should be added the 
records from Washington (Banner, 1948, pt. I, p. 381). 


Pages 137, 244.—Caesaromysides liguriae Colosi 


This genus should be cancelled as an immature stage of Caesaromysts; 
the worth of the species could not be determined because of the 
immaturity of the specimen described (Banner, 1948, pt. I, p. 389). 

To Tattersall’s account of American mysids, Caesaromysis vanclever 
Banner (loc. cit.,) should be added here, with its records from off 
Oregon to off Alaska. 


Page 160.—Inusitatomysis serrata Tattersall 


To this species should be added the reference Jnusitatomysis sp. 
Banner (1948, pt. II, p. 67), which is apparently the same species as 
Tattersall’s J. serrata. One major difference between the two sets of 
specimens is in the depth of the terminal cleft of the telson, which is 
one-eighth the length of the telson in Tattersall’s specimens and one- 
third the length in mine; however, this may well be a growth difference 
and the two groups of specimens are probably the same species. 

If they are the same, then the records of Inusitatomysis sp. from off 
British Columbia should be added to Tattersall’s record from the 
Bering Sea. 


Page 165.—Mysis oculata (Fabricius) 


To the list of synonyms of this species should be added Pugetomysis 
litoralis Banner (1948, pt. II, p. 104), and the range of the species in 
the Pacific should be extended to include the waters of Washington.° 


6 TI plan to discuss this change in a forthcoming publication. 


582 sv 1+: .PROCEEDINGS. OF THE NATIONAL MUSEUM: ©; vot. 103 


Page.181.—-Neomysis rayii (Murdoch) 

Tattersall and Banner arrived independently at the same conclusion 
that N. franciscorum Holmes is a synonym of N. rayti. The records 
given by Banner (1948, pt. II, p. 78) augment those given by Tattersall. 


Page 187. —Neomysis mercedis Holmes 
Under this species should be added the observations that it pene- 
trates and lives in fresh-water lakes, such as Lake Washington near 
Seattle (Banner, 1948, pt. II, p. 75).’ 
Page 192.—Neomysis kadiakensis Ortmann 
Additional records of this species from Washington and British 
Columbia have been given (Banner, 1948, pt. II, p. 82). 


Pages .208,'248.—Acanthomysis sculpta (Tattersall) 

Observations have shown that this species exhibits great variation 
on many points, especially in the sculpturing of the abdomen (Banner, 
1948, pt. II, pp. 97-101). The records have extended the range of 
the species from Washington to southern California. 


Page 215.—Acanthomysis macropsis (Tattersall) 


The known range of this species has been extended to include the 
inland waters of Washington (Banner, 1948, pt. I, p. 91). 


Page 217.—Acanthomysis pseudomacropsis (Tattersall) 

Tattersall, in an earlier publication (1933, p. 94) and again in this 
publication, states that he believes the exopod of the fourth pleopods of 
the male to be of three articles; this appendage was found to be com- 
posed of a single article (Banner, 1948, pt. II. p. 89). In addition, the 
records cited have extended the range southward into Washington and 
also give another Alaskan record. 


Page 218. 

To the genus Acanthomysis should be added two new species 
described from the northwestern coast of America, A. nephrophthalma 
Banner and A. davisi Banner (1948, pt. II, pp. 93, 95), and an unnamed 
specimen from Alaska, Acanthomysis sp. Banivios (1948, pt. II, p. 101). 
Pages 220, 249.—Proneomysis wailesi Tattersall 

en an additional record for this species from Washington a new 
location has been given (Banner, 1948, pt. II, p. 104). 
Pages 242 or 250. 

To Tattersall’s list of genera and species occurring in the waters of 
the Americas should be added Mysidella americana Banner (1948, 
pelh p. 109), a member of the last subfamily of the Mysidae, the 
Mysidellinae, which was described from British Columbia and Shick 
is the only record of the subfamily from American waters. 


7I plan to discuss the validity of this species in a forthcoming publication. 


SUPPLEMENT TO TATTERSALL’S MYSIDACEA—BANNER 583 


References 


BActsco, Minar 

1941a. Sur une petite collection de Mysidacés provenant de Villefranche- 
sur-Mer (Méditerranée). Arch. Zool. Exp. et Gén, vol. 81, pt. 
4, pp. 164-172. 

1941b. Les Mysidacés des eaux méditerranéenes de la France (spéciale- 
ment de Banyuls) et des eaux de Monaco. Bull. Inst. Océanogr. 
Monaco, No. 795, pp. 1-46. 

BANNER, ALBERT H. 

1948. <A taxonomic study of the Mysidacea and Euphausiacea (Crustacea) 
of the Northeastern Pacific. Part I, Mysidacea, from family 
Lophogastridae through tribe Erythropini; Trans. Roy. Canadian 
Inst., vol. 26, pp. 345-397, pls. 1-9. Part II, Mysidacea, from 
tribe Mysini through subfamily Mysidellinae; Trans. Roy. Cana- 
dian Inst., vol. 27, pp. 65-125, pls. 1-7. 

Faces, Louis 

1941. Mysidacea. Lophogastrida—I. Dana Report, No. 19, 52 pp., 51 
figs. 

1942. Mysidacea. Lophogastrida—II. Dana Report, No. 23, 67 pp., 42 
figs. 

NovuveE., HENRI 

1942a. Sur la systématique des espéces du genre Hucopia Dana, 1852 (Crust. 
Mysidacea). Bull. Inst. Océanogr. Monaco, No. 818, 8 pp. 

1942b. Diagnoses préliminaires de Mysidacés nouveaux provenant des 
Campagnes du Prince Albert I¢*de Monaco. Bull. Inst. Océanogr. 
Monaco, No. 831, 12 pp. 

1948. Mysidacés provenant des Campagnes du Prince Albert I*t de Monaco. 
In Résultats des Campagnes Scientifiques . . . par AlbertI... 
de Monaco, vol. 105, pp. 1-128, pls. 1-5. 

TATTERSALL, WALTER M. 

1933. Euphausiacea and Mysidacea from western Canada. Contr. Cana- 
dian Biol. Fish., new. ser., vol. 8, No. 15, pp. 181-205. 

1951. A review of the Mysidacea of the United States National Museum. 
U.S. Nat. Mus. Bull. 201, x + 292 pp., 108 figs. 

TATTERSALL, WALTER M., and TaTTERSALL, OLIVE 8. 
1951. The British Mysidacea. Ray Society Publ. No. 136, pp. viii + 460. 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington : 1955 No. 3335 


ON THE ANATOMY AND RELATIONSHIPS OF GLOSSY 
CUCKOOS OF THE GENERA CHRYSOCOCCYX, 
LAMPROMORPHA, AND CHALCITES 


By AnpREw J. BERGER’ 


The relationships of the glossy cuckoos have long been in question. 
Sharpe (1873, p. 579) commented that he could not “find any charac- 
ter in the genus Chrysococcyz to justify its separation from Cuculus, 
beyond the metallic plumage...” Shelley (1891, pp. 280, 288) 
recognized as distinct the genera Chrysococcyx and Chalcococcyx 
(= Chalcites), but placed Lampromorpha in synonymy with Chryso- 
coccyx. Bannerman (1922, pp. 4138-420; 1933, pp. 111-120), how- 
ever, recognized both Lampromorpha and Chrysococcyx. Chapin 
(1939, pp. 197-201), Peters (1940, p. 29), and Friedmann (1948, p. 
115) again placed Lampromorpha in synonymy with Chrysococcyz. 
Delacour (1951, p. 19) and Friedmann (in litt.) further suggest that 
the genus Chalcites should be united with Chrysococcyx, which would 
place all species of this complex in the genus Chrysococcyzx. 

Little has been written on the internal anatomy of Chrysococcyx 
and, so far as I know, there has been no published description of the 
internal anatomy of Lampromorpha or Chalcites. 

I am indebted to the following for supplying alcoholic and skeletal 
material: Dr. Dean Amadon, American Museum of Natural History; 
Dr. Herbert Friedmann, United States National Museum; and Mr. 
John G. Williams, the Coryndon Museum, Nairobi, East Africa. I 
had the following spirit material for study: Chrysococcyx cupreus 
(8), Lampromorpha caprius (2), L. klaas (1), and Chalcites 


1 Department of Anatomy, University of Michigan Medical School, Ann Arbor, Michigan. 
297268—55 585 


586 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


lucidus (2). Skeletal material was available as follows: Chrysococeyx 
cupreus (3), Lampromorpha caprius (2), and L. klaas (1). Certain 
information was also obtained from the skeletons of the alcoholic 
specimens. For reading the manuscript and for offering suggestions 
I am indebted to Dr. Amadon, Dr. Jean Delacour, Los Angeles 
County Museum, California, Dr. Friedmann, and Dr. Josselyn Van 
Tyne, University of Michigan Museum of Zoology. 


Pterylosis 


Beddard (1885, p. 187), basing his comments on manuscript notes 
of Garrod and Forbes, grouped Chrysococcyx with those cuckoos 
which have the “ventral tract [abdominal] of both sides single,” but 
this arrangement is, in fact, not found in Chrysococeyx. Miller (1924, 
p- 3830) reported that the nestling of Chrysococcyx lacks natal down. 
Nothing else seems to have been published on the pterylosis of the 
species discussed in this paper. 

In general the pterylosis of the four species is similar. In each 
species there are 10 primaries and 10 secondaries, though secondaries 
8, 9, and 10 (outermost counted as first) are progressively smaller. 
The fifth secondary is present, i. e., the wing is eutaxic, or quintocu- 
bital. There are four alula quills, the outermost being the longest. 
The carpal remex and its covert are present. There are 10 rectrices 
and 10 upper tail coverts. The oil gland is nude; a single, median 
feather at the base of the gland passes caudad, covering it. The eye- 
lashes are featherlike rather than hairlike as in Coua caerulea. The 
tarsi are what Sharpe (1873, p. 579) spoke of as “accipitrine,” i. e., 
feather tracts descend for a distance of about 6 mm. on the tarsometa- 
tarsus; the lateral tract shows the best development. 

In the capital region, the median frontal apterium is wanting. A 
small supraorbital apterium is present, but it is separated from the 
eyelashes by a single row of feathers. The dorsal cervical feather 
tract is continuous with the interscapular tract. A wide lateral 
cervical apterium is present. A long, narrow, median dorsal apterium 
extends through the dorsal region and the anterior half of the pelvic 
region. There is a single, medium pelvic feather tract. 

Bilateral submalar apteria are present; these are widest in Chryso- 
coccyx and narrowest in Lampromorpha caprius. 

The abdominal branches of the above-mentioned genera differ from 
those shown for Cacomantis, Piaya (Beddard, 1885), and Cuculus 
canorus (Lowe, 1948, p. 493) in that two separate tracts form at the 
posterior margin of the sternum. An inner abdominal tract extends 
to the anus; this tract is two feathers wide at the beginning, but is re- 
duced to one row about two-thirds the distance to the anus. The outer 
abdominal tract turns inward to meet the inner abdominal tract about 
midway to the anus. 


ANATOMY AND RELATIONSHIP OF GLOSSY CUCKOOS—BERGER 587 


Osteology 


All four species have 14 cervical vertebrae. There are two pairs 
of cervicodorsal ribs and, usually, only the last such rib possesses an 
uncinate process, though one specimen of Chrysococcyx cupreus has 
uncinate processes on both left cervicodorsal ribs, one specimen of 
Lampromorpha caprius possesses these processes on both right ribs, 
and an alcoholic specimen of Chalcites lucidus has uncinate processes 
on all four ribs. The atlas is perforated by the odontoid process, ex- 
cept in a strongly bleached skeleton of Lampromorpha klaas, where 
the atlas appears to be notched. 

With one exception, each specimen has four dorsal vertebrae, four 
dorsal ribs, and one thoracic rib. In each case, the thoracic rib articu- 
lates with the synsacrum dorsally and fuses with the last dorsal rib 
ventrally. One alcoholic specimen of Chrysococcyx cupreus, however, 
has five dorsal vertebrae and five dorsal, or true, ribs. Of the latter, 
four articulate with the sternum, while the fifth fuses ventrally with 
the fourth rib; there is no thoracic rib. Of ten cuculine genera I have 
thus far investigated, this is the first example I have seen of intra- 
specific variation in the number of dorsal vertebrae. The number 
of ribs articulating directly with the sternum, on the other hand, is 
far more variable. Four, apparently, is the “normal” number of 
sternal ribs in these species. In one specimen of Chrysococcyx cupreus 
and two specimens of Lampromorpha caprius, however, only three of 
the four dorsal ribs articulate directly with the sternum; and in an 
alcoholic specimen of L. klaas three ribs articulate with the sternum 
on the left side, whereas four do so on the right. 

Apparently in younger individuals the general pattern of the pos- 
terior emargination of the sternum is double-notched ; it does not seem 
likely that the bone would be resorbed to form fenestrae in older 
individuals. In half of the specimens, the sternum is single-notched 
and bilateral fenestrae are present. In some cases, these fenestrae are ° 
bounded posteriorly by a bony bar only 1 mm. in width. 

The external nares are oval to rounded and their borders are con- 
spicuously swollen. On the floor of the narial cavity is a well-de- 
veloped tubercle which has a small bony core; the latter is readily 
discernible in the skeleton. 

A pectineal process is not developed. 


Wing Myology 


In all four species the following wing muscles are absent: pro- 
scapulohumeralis brevis, abductor indicis brevis, abductor digiti II, 
and the biceps slip to the tendon of M. tensor patagii longus. Gadow’s 
pars propatagialis musculi cucullaris does not insert on the tendon of 
M. tensor patagii longus. 


588 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


P. pro. 


Pect. tend. 


Ext. meta. 


Figure 69.—Chrysococcyx cupreus. Anconal view of left arm and forearm to show the 
structure and relationships of M. tensor patagii brevis. Explanation of symbols: Del., 
deltoideus major; Ext. dig., extensor digitorum communis; Ext. meta., extensor meta- 
carpi radialis; P. pro., pars propatagialis, pectoralis superficialis; Pect. tend., accessory 
tendon of pectoralis superficialis; Tpb., tensor patagii brevis; Tpl., tensor patagii longus. 


The structure of M. tensor patagii brevis (fig. 69) is similar in the 
three genera and exhibits some differences from that found in other 
cuckoos studied. In Lampromorpha caprius the belly is 14 mm. long 
and 5 mm. in maximum width. The distal end of its fleshy belly has 
no relationship to the tendon of M. tensor patagii longus. A strong flat 
tendon (0.5 mm. wide) arises from the dense fascial envelope which 
surrounds the insertion of M. pectoralis superficialis, and fuses with 
the tendon of M. tensor patagii brevis. About 5 mm. distal to the 
ectepicondylar process of the humerus, part of the tendon of M. tensor 
patagii brevis fuses with the superficial surface of M. extensor meta- 
carpi radialis. From this region of fusion, a strong tendon passes 
proximad to attach to the ectepicondylar process; distal to the area 
of fusion, the tendon expands and gives origin on both its superficial 
and deep surfaces to fleshy fibers of M. extensor metacarpi radialis. 
Posterior to the area of fusion, the tendon of M. tensor patagii brevis 
bifurcates: one branch fuses with the superficial surface of M. exten- 
sor digitorum communis and attaches to the humerus with that 
muscle; the second branch fans out, extends nearly the entire length 
of the ulna, attaches to the bases of the secondaries, and fuses with the 
antibrachial fascia. This muscle shows the same relationships in the 
other species, but measurements are slightly smaller in them. In its 
insertion, M. tensor patagii brevis is similar to that in Coccyzus 
erythropthalmus. 


ANATOMY AND RELATIONSHIP OF GLOSSY CUCKOOS—BERGER 589 


Ext. meta. tend. 


T. p. long. 


Flex. c. brev. 


Acc. flex. 


1 cm. 
fener ee ee ey 


Figure 70.—Chrysococcyx cupreus. Palmar view of part of left wing to show relationships 
of the accessory flexor muscle of the pollex. Explanation of symbols: Abd. p., abductor 
pollicis; Acc. flex., accessory flexor muscle; Ext. meta. tend., tendon of extensor meta- 
carpi radialis; Flex. c. brev., flexor carpi ulnaris brevis; T. p. long., tendon of tensor 
patagii longus. 


Although M. tensor patagii longus (figs. 69, 70) has a common 
origin with M. tensor patagii brevis from the superomedial surface 
of the furculum, the two bellies are incompletely fused distally. In 
Lampromorpha caprius, M. pectoralis, pars propatagialis, is a well- 
developed, fleshy fasciculus 14 mm. in length and 5 mm. wide at its 
base on the furculum. The belly tapers to a tendon whose direct con- 
tinuation is the tendon of M. tensor patagii longus. The belly of the 
latter muscle (about 8 mm. long and 2 mm. wide) fuses with M. pec- 
toralis, pars propatagialis, just before the formation of the common 
tendon. The tendon runs distad in the leading edge of the propa- 
tagium and bifurcates near the distal end of the radius. The main 
tendon fuses with the fascia of the manus and has a minor attachment 
to the extensor process. The smaller tendon serves as the origin for 
a small (4mm. long) fleshy belly (fig. 70) which inserts on the postero- 
basal corner of the pollex superficial to the insertion of M. flexor pol- 
licis, which, although present, is poorly developed. CArysococcyx and 
Chalcites exhibit this same structure, which I have found neither in 
other cuckoos nor in literature on other birds. 

M. coracobrachialis anterior is similar in the four species and, as in 
Coccyzus erythropthalmus, it is poorly developed. The belly is sur- 
rounded by a dense fascia and, in fact, the muscle almost seems to be 


590 PROCEEDINGS OF THE NATIONAL MUSEUM \VOL. 103 


imbedded in the joint capsule. The belly does not fan out at its 
insertion. 

M. expansor secundariorum is a single muscle with two tendons of 
origin: ascapular tendon and a humeral tendon. 

M. scapulotriceps does not have an accessory attachment to the 
humerus. 

M. subcoracoideus has two heads, one arises from the furculum and 
the other arises from the coracoclavicular membrane as in other 
cuckoos. 

M. deltoideus minor is present and arises inside the triosseal canal 
from the inferior furcular process. 

In all four species, the tendon of M. latissimus dorsi, pars posticus, 
inserts posterior to and in contact with the tendon of pars anticus. 
In Chalcites, Chrysococcyx, and Lampromorpha klaas the area of in- 
sertion of pars posticus begins about .5 mm. proximal to the insertion 
of pars anticus, whereas in Lampromorpha caprius the insertion of 
pars posticus begins about .5 mm. distal to the uppermost fibers of 
insertion of pars anticus. A dermal component is present, but is very 
small, and I was unable to demonstrate it in every specimen. 

In each species, M. pronator longus is a relatively short muscle, ex- 
tending distad the same distance as M. pronator brevis, i. e., about two- 
thirds the length of the radius. 

M. flexor digitorum profundus arises on both sides of the area of 
insertion of M. brachialis. 

M. flexor carpi ulnaris brevis (fig. 70) shows the best development 
yet seen in cuckoos. The fleshy belly is relatively and absolutely 
longer (14 mm. in Chrysococcyx and Lampromorpha caprius) than 
in Coua, Geococcyzx, etc., and the belly is located more proximally on 
the ulna. 

M. flexor metacarpi brevis is well developed in all four species. The 
belly is triangular in shape; it arises fleshy from the base of the 
carpometacarpus; it inserts on the tendon of M. extensor indicis 
longus. 

M. extensor pollicis brevis exhibits minor variation. In Chryso- 
coccyx cupreus none of its fibers appear to arise from the tendon of M. 
extensor pollicis longus. In ZLampromorpha caprius there are two 
heads as in Crotophaga sulcirostris. In Chalcites the structure is in- 
termediate between that found in Crotophaga and in Coccyzus, and a 
few fibers arise from the tendon of M. extensor pollicis longus. 

M. abductor pollicis is a small muscle with a maximum length of 
6mm. I found two bellies to this muscle in Chrysococcyx cupreus, 
but I was unable to determine to my satisfaction in preserved material 
that there are two parts in Lampromorpha or Chalcites. The origin 
and insertion of this muscle are in each case, however, typical for 
cuckoos. 


ANATOMY AND RELATIONSHIP OF GLOSSY CUCKOOS—BERGER 591 
Leg Myology 


The muscle formula in the three genera is AX YAm. The following 
muscles are absent: pars iliofemoralis of the piriformis, iliotrochan- 
tericus medius, gluteus medius et minimus, iliacus, adductor digiti IV, 
extensor brevis digiti III, and, apparently, adductor digiti Il A 
vinculum is present between the tendons of Mm. fiexor digitorum 
longus and flexor hallucis longus, but a vinculum is wanting between 
the tendons of Mm. flexor perforatus digiti III and flexor perforans et 
perforatus digiti III. The tendon of insertion of M. flexor perforatus 
digiti If forms a sheath for the tendon of M. flexor perforans et 
perforatus digiti II, but the latter tendon does not ensheath the 
tendon of M. fiexor digitorum longus which supplies digit IL. The 
tendon of M. flexor perforatus digiti IV does not ensheath the tendon 
of M. flexor digitorum longus which supplies digit IV. 

In Lampromorpha caprius, M. iliotibialis (fig. 71) arises semiten- 
dinous from the entire anterior iliac crest, the anterior iliac process, 
and from approximately the anterior half of the posterior iliac crest. 
The muscle is composed of anterior and posterior fleshy portions 
separated by a central aponeurosis, which distally is intimately fused 
with the underlying M. femorotibialis externus. The central aponeu- 
rosis is best developed distally, where it is 13 mm. long and 3.5 mm. in 
maximum width; proximally, this band is only 1 mm. wide. In this 
central area, however, the muscle is completely aponeurotic from 
origin to insertion. Except for its smaller size, M. iliotibialis exhibits 
the same structure in Chalcites lucidus. In Chrysococcyx cupreus 
the distal aponeurotic portion is the same shape and size as in 
Lampromorpha, but it is separated from the aponeurosis of origin 
by a central fleshy portion 5 mm. long; for this short distance, there- 
fore, the belly is entirely fleshy from anterior to posterior border. 
In each of the genera, the insertion of M. iliotibialis is typical and 
its belly conceals only the anterior and proximal half of M. biceps 
femoris. 

M. semitendinosus (fig. 71) is similar in the three genera. It arises 
from the posterior third of the posterior iliac crest and from the 
transverse process of the first free caudal vertebra. Its insertion is 
typical. 

M. gastrocnemius is composed of three main parts: pars externa, 
pars media, and pars interna. Pars externa arises by two heads, as 
in Coccyzus. Pars media is very small: 5, 7, and 8 mm. long, respec- 
tively, in Chalcites, Lampromorpha, and Chrysococcyx. The fleshy 
portions of pars externa and pars interna are nowhere in contact; 
the posterior aspect of the complex is a thin, tendinous sheet, which 
connects the fleshy bellies. 


592 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


M. peroneus longus is poorly developed, resembling the structure 
of that muscle in Coccyzus. Fleshy fibers begin some distance (7 to 
10 mm.) distal to the rotular crest, and the belly does not conceal any 
part of M. tibialis anticus. 

M. plantaris is a long, thin muscle whose belly extends about two- 
thirds the distance down the tibiotarsus. 

In Chrysococcyz cupreus, M. extensor proprius digiti IIT is a long 
thin muscle which begins just distal to the proximal articular surface 
of the tarsometatarsus and extends to the distal end of that bone. The 
belly does not increase much distad, as it does in Coccyzus. The muscle 
is similar in origin in Chalcites lucidus, but the fleshy fibers end 5.5 
mm. proximal to the insertion. In Lampromorpha caprius, the muscle 
arises at the level of insertion of M. tibialis anticus. 

M. lumbricalis exhibits its best development in Lampromorpha 
and its poorest development in Chalcites; in the latter genus it is 
limited to the distal half of the tarsometatarsus. 

M. obturator internus, similar in the three genera, is triangular in 
shape and has only a small fasciculus arising from inside the pelvis. 


Syrinx 


The syrinx is tracheobronchial. There is one pair of intrinsic 
syringeal muscles. In Chrysococcyx cupreus and Lampromorpha 
klaas these muscles insert on the fourth bronchial semiring. This ring 
is not much wider than adjacent rings, but it has the largest diameter 
of any in the bronchial series. The structure is similar in L. caprius 
and Chalcites lucidus except that the muscles insert on the third semi- 
ring. However, not more than two syringes were suitable for study 
in any one species so I cannot say that the point of insertion is constant. 


Viscera 


In each specimen the right lobe of the liver is much larger than the 
left lobe. A gall bladder is not evident. The pancreas is bilobed. 
One can obtain only an approximate total-length measurement of the 
intestine. Its total length in millimeters in two specimens of Lampro- 
morpha caprius is 220 and 240; in two specimens of Chalcites lucidus, 
160 and 180; and in two specimens of Chrysococcyx cupreus, 170 and 
260. Only insect remains were found in the stomachs. <A peculiar 
feature is the presence of a single caecum, about 4 mm. long in Chaleites 
and 5 mm. long in Lampromorpha and Chrysococcy#. 


Discussion 


The genera Chrysococcyx and Lampromorpha are restricted to 
Africa. Chalcites is found in Asia, Australia, and New Zealand. 


ANATOMY AND RELATIONSHIP OF GLOSSY CUCKOOS—BERGER 593 


Dey 


) nal 
\\f | Bic. fem. 
Aponeurotic 17 ! 
1 cm. 
fered 


Ficure 71.—Lampromorpha caprius. Superficial muscles of the left thigh (lateral view). 
Explanation of symbols: Bic. fem., biceps femoris; Il, tib., iliotibialis; Sar., sartorius; 
Semim., semimembranosus; Semit., semitendinosus. 


Except for the rare yellow-throated cuckoo (Chrysococcyx flavigu- 
laris), which has not been studied in the field, it is known that all of 
the glossy cuckoos are parasitic on other birds. A large number of 
hosts have been recorded, but for the African species the most im- 
portant hosts seem to be members of the following families: Ploceidae, 
Nectariniidae, Sylviidae, Muscicapidae, and Motacillidae. Common 
hosts of the bronze cuckoos (Chalcites) in Australia are certain thorn- 
bills, wrens, and scrub-wrens of the family Sylviidae (Serventy and 
Whittell, 1948, p. 238). For other species of Chalcites in India, Baker 
(1984, pp. 853-354) reports the most common hosts to be sunbirds 
and spider-hunters (Nectariniidae). 

Friedmann (1948, pp. 115-116) remarked that: “The eggs of the 
glossy cuckoos show far more variation in color and markings than do 
those of any of the other cuckoos of Africa.” The eggs of Klaas’s 
cuckoo (Lampromorpha klaas), for example, have “the ground 
color . . . white, pale greenish blue, or pale blue, and either unspotted 
or quite abundantly flecked with brick red, brown, or purplish gray,” 
(ibid., p. 186). According to Barrett (1945, p. 149), “five kinds of 
bronze cuckoos occur in Australia, and the eggs of four of them are 
bronzy-green in colour, while that of the fifth, the narrow-billed bronze 


594 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


cuckoo (Chalcites basalis) is pinkish, with a uniform sprinkling of 
tiny reddish spots.” Serventy and Whittell (1948, p. 238), in speaking 
of Chalcites lucidus, state: “The egg is pale olive in colour, without 
any markings, but this colour can be removed with water, after which 
the shell is found to be light bluish in colour.” 

Friedmann (1948, p. 115; 1949, p. 519) has called attention to a 
behavior pattern “unique” among parasitic birds in that the “habit 
of feeding of fledged young by adult parasitic cuckoos is known... 
only in some four species of the small glossy cuckoos of the Chalcites- 
Chrysococcyx group.” However, Moreau (1949, p. 537) cited two 
references indicating that adults of Cuculus occasionally feed young 
cuckoos. 

“The African glossy cuckoos also have a courtship feeding behavior,” 
which “is otherwise known, in cuckoos, only in African forms of 
Clamator and Cuculus” (Friedmann, 1948, p. 115). Since this state- 
ment was published, however, Watson and Bull (1950, p. 226) have 
reported observations of Chalcites lucidus which gave them the “im- 
pression . . . that some form of communal display with courtship 
feeding was taking place.” These authors do not agree with Fried- 
mann (1949, p. 517) that feeding of young by adults has been proved 
for the New Zealand shining cuckoo. 

That the species of glossy cuckoos here discussed are closely related, 
biologically and anatomically, is evident from an analysis of the 
data presented. The behavior patterns of courtship feeding and feed- 
ing of fledged young indicate close relationship. Too little is known 
about the detailed life history of these birds to justify further com- 
parison. In morphological characters, also, these species exhibit a 
general uniformity of structure. For the most part, differences in 
muscles, for example, can be ascribed only to size differences of the 
species. 

It seems certain that unsatisfactory characters have been used by 
some authors to separate Chrysococcyx and Lampromorpha. Jackson 
(1938, p. 500) used shape and length of tail to separate them: Lam- 
promorpha “has the tail much shorter and slightly rounded, but not 
graduated.” Priest (1934, p. 227) separated them on the basis of 
sexual dimorphism, but this surely was a misprint in his key. In 
establishing the genus “A damatornis” for klaas, Roberts (1922, p. 219) 
stated only that klaas “is much smaller than cupreus.” 

Little has been written on the locomotor habits of these species. 
Yet, of seven differences in anatomical structure which might be useful 
for determining relationship, six seem definitely to be correlated to 
functional phenomena. These involve primarily the extent of origin 
or insertion and relative length of belly of the following muscles: 
latissimus dorsi, iliotibialis, extensor proprius digiti III, abductor 


ANATOMY AND RELATIONSHIP OF GLOSSY CUCKOOS—BERGER 595 


pollicis, flexor hallucis longus, and extensor hallucis longus. It must 
be emphasized, however, that all differences in structure here dis- 
cussed are minor in degree. 

An analysis of M. iliotibialis will serve to illustrate the minor dif- 
ferences encountered and to cast doubt on the validity of them as a 
basis for establishing separate genera. The structure of this muscle 
has been discussed in some detail above (p. 591). The complete apo- 
neurotic central portion of M. iliotibialis in Lampromorpha caprius 
is most like the structure of that muscle in Coccyzus. In each of eight 
other genera (Crotophaga, Geococcyx, Coua, etc.) the aponeurotic 
portion is limited to the distal half or two-thirds and there is a well- 
developed fleshy belly located proximal to this aponeurosis. Only in 
Chrysococcyx cupreus and Lampromorpha klaas, where the middle 
fleshy portion is very small, does the structure of the muscle 
represent an intermediate stage. The locomotor pattern seems to be 
similar in these species (Friedmann (1948, pp. 129, 150, 158) implies 
this as regards the manner of flight), so one might place some 
weight on this character as indicating degree of relationship. Also, 
the place of insertion of the syringeal muscles, if proven constant, 
would indicate relationship as follows: caprius with lucidus (insertion 
on the third bronchial ring) ; cwpreus with klaas (insertion on the 
fourth ring). Minor differences in other muscles, however, indicate 
a closer relationship between lucidus and cupreus than between the 
former and caprius, so that it seems unlikely that the structure of 
M. iliotibialis, in this instance, is a valid character for judging rela- 
tionship. In anatomical features klaas resembles cupreus more than 
it does caprius. Van Someren (1925, pp. 660-662) came to the same 
conclusion after a study of feather structure; he stated that klaas 
should either be placed with cupreus in the genus Chrysococcyx or 
else in an intermediate position, but that 4/aas could not be placed with 
caprius if only two genera were to be recognized. Both van Someren 
and Friedmann (1948, pp. 150, 151) felt that cupreus, caprius, and 
klaas are congeneric. 

In reviewing the evidence presented here, I must conclude that all 
of the glossy cuckoos are congeneric. Following the law of priority, 
all the species of this complex must be placed in the genus Chryso- 
coccyx. Anatomical similarities further suggest that cupreus and 
klaas are more closely related to each other than to either caprius or 
lucidus. Though Jucidus possesses certain anatomical features ex- 
hibited by caprius (especially the structure of M. iliotibialis and the 
place of insertion of the syringeal muscles), in most respects luctdus 
is like cupreus. Therefore, caprius is most unlike the other species 
of this genus. 


596 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


References 


Baker, EB. C. Stuart 
1934. The nidification of birds of the Indian Empire, vol. 3, 568 pp., 8 pls. 
BANNERMAN, Davip A. 
1922. On the emerald and golden cuckoos of Africa. Nov. Zool., vol. 29, 
No. 2, pp. 413-420. 
1933. The birds of tropical West Africa, vol. 3, 487 pp., 144 figs., 12 pls., map. 
BaRRETT, CHARLES 
1945. Australian bird life, 239 pp., 46 pls. 
BEpDARD, FRANK BH. 
1885. On the structural characters and classification of the cuckoos. Proc. 
Zool. Soc. London (1885), pp. 168-187. 
CHAPIN, JAMES P. 
1939. The birds of the Belgian Congo. II. Bull. Amer. Mus. Nat. Hist., vol. 
75, 632 pp., 38 figs., 21 pls. 
DELACOUR, JEAN 
1951. Commentaires, modifications et additions a la liste des oiseaux de 
l’Indochine Frangaise. L’Oiseau Rev. Francaise d’Ornith., vol. 21, 
pp. 1-32. 
FRIEDMANN, HERBERT 
1948. The parasitic cuckoos of Africa. Washington Acad. Sci. Monogr. 
No. 1, 204 pp., 10 pls. 
1949. Additional data on African parasitic cuckoos. Ibis, vol. 91, No. 3, 
pp. 514-519. 
JACKSON, FREDERICK J. 
1938. The birds of Kenya Colony and the Uganda Protectorate, vol. 1, 542 pp., 
115 figs., 10 pls. 
Lowe, Prrcy R. 
1948. Some notes on the anatomical differences obtaining between the Cucul- 
idae and the Musophagidae, with special reference to the specializa- 
tion of the oesophagus in Cuculus canorus Linnaeus. Ibis, vol. 85 
(October 1948), pp. 490-515. 
MATHEWS, GREGORY M. 
1918. The birds of Australia, vol. 7, pts. 1-4, pp. 1-384, pls. 325-362. 
1919. The birds of Australia, vol. 7, pt. 5, pp. 385-499, pls. 363-479. 
MILLER, W. DreWI1TT 
1924. Further notes on ptilosis. Bull. Amer. Mus. Nat. Hist., vol. 50, art. 


5, pp. 305-331. 
Moreau, R. E. 
1949. Friedmann on African cuckoos (special review). Ibis, vol. 91, pp. 
529-537. 


PETERS, JAMES L. 
1940. Check-list of birds of the world, vol. 4, 291 pp. 
Prirest, Ceciu D. 
1934. The birds of Southern Rhodesia, vol. 2, 553 pp., 172 figs., 10 pls. 
Rogerts, AUSTIN 
1922. Review of the nomenclature of South African birds. Ann. Transvaal 
Mus., vol. 8, pp. 187-272. 
SERVENTY, D. L., and WHITTELL, H. M. 
1948. A handbook of the birds of western Australia, 365 pp., 32 figs., 2 pls. 
SHARPE, R. BOWDLER 
1873. On the Cuculidae of the Ethiopian region. Proc. Zool. Soc. London 
(1878), pp. 578-624. 


ANATOMY AND RELATIONSHIP OF GLOSSY CUCKOOS—BERGER 597 


SHELLEY, G. E. 
1891. Picariae—Scansores and Coccyges, in British Museum catalogue of 
birds, vol. 19, 484 pp., 13 pls. 
VAN SOMEREN, V. G. L. 
1925. What is the true relationship of Klaas’s cuckoo to Ohrysococcyz 
cupreus and Lampromorpha caprius? Ibis, ser. 12, vol. 1, pp. 660— 
662, pls. 22, 23. 
Watson, J.S., and Butt, P. C. 
1950. Communal display of the shining cuckoo. New Zealand Bird Netes, 
vol. 3, p. 226. 


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SMITHSONIAN INSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1955 No. 3336 


SUPPLEMENT AND CORRECTIONS TO J. A. HYSLOP’S 
GENOTYPES OF THE ELATERID BEETLES OF THE 
WORLD' 


By Ross H. Arnett, Jr.? 


The publication in 1921 of J. A. Hyslop’s “Genotypes of the Elaterid 
Beetles of the World” (Proc. U. 8. Nat. Mus., vol. 58, No. 2353, 
pp. 621-680) opened the road to a stabilized nomenclature in the 
family Elateridae. The impetus of this work resulted in a great many 
studies in the generic classification of this moderately large family. 
Although many of the generic names employed up to that time were 
shown to be incorrect, a great stride was made in providing a means 
for the stabilization of the names in this group, because, as I believe 
has been shown repeatedly, it is only after the genotypes have been 
correctly determined that generic assignments can be made. 

However, during the 34 years since the publication of this work, a 
considerable number of new names have been proposed, and a fair 
number of inaccuracies have been discovered. Many of these errors 
have been pointed out by Méquignon and Lane in the following 
papers: 

Méaquienon, A. 

1930. Notes synonymiques sur les Elatérides . . . Bull. Soc. Ent. France 
(1930), pp. 91-96. 

LANE, M. C. 

1948. Some generic corrections in the Elateridae—I. Proc. Ent. Soc. Wash- 
ington, vol. 50. pp. 179-182. 

1948, Some generic corrections in the Elateridae—II. Proc. Ent. Soc. Wash- 
ington, vol. 50, pp. 221-223. 

1949. Some generic corrections in the Elateridae—III. Proc. Ent. Soc. 
Washington, vol. 51, pp. 152-153. 

1953. Some generic corrections in the Elateridae—IV. Proc. Ent. Soc. 
Washington, vol. 55, pp. 86-89. 


1Proc. U. 8S. Nat. Mus., vol. 58, No. 2353, pp. 621-680, 1921. 
?Formerly of Entomology Research Branch, Agriculture Research Service, U. S. Department of 
Agriculture; now Head, Department of Biology, St. John Fisher College, Rochester, N. Y. 


309150—55——1 599 


600 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


The following pages are intended as a supplement to Hyslop’s paper 
to bring together these corrections and additions, as well as to add 
further corrections. There are a number of minor typographic mis- 
takes in Hyslop’s paper which I have not listed below, and in a few 
cases he gives incorrect dates which are obvious to anyone using the 
catalog (i. e., Linnaeus ‘‘1857,”’ etc.) and which have not been corrected 
here. With these exceptions, it is thought that this list is correct to 
and including part of 1952. 


New names 


As a result of this study, three new names are necessary to replace 
others which are shown to be unavailable. 
Malloea new subgenus. 

TypE: Ctenicera (Malloea) sjaelandica (Miiller) (=Elater sjae- 
landicus Miiller), Fauna Insectorum Fridrichsdalina, p. 21, 
1764. (Here designated.) 

Nore: This subgenus of Ctenicera includes those species grouped 
under the subgeneric name Actenicerus Kiesenwetter in the Cole- 
opterorum Catalogus (Schenkling, 1927, pt. 88, pp. 368-370, 1927) 
and the species described by Kiesenwetter under the name Actenicerus- 
The name Actenicerus Kiesenwetter, 1857 (Naturgeschichte der 
Insecten Deutschlands, vol. 4, p. 285), is not available because it is a 
junior synonym of Prosternon. Corymbites tessellatus Germar, 
(Zeitschr. Ent., vol. 4, p. 62, 1843), which Kiesenwetter gives as the 
only included species in his subgenus Actenicerus, is referred back to 
Elater tessellatus Linnaeus (Systema Naturae, ed. 10, p. 406, 1758), 
a species which is now included in the genus Prosternon Latreille, 1834. 
The Elater tessellatus of Fabricius (Systema Entomologiae, p. 211, 
1775), which Kiesenwetter redescribed and misdetermined as 
Corymbites tessellatus Germar, is Elater sjaclandicus Miller, 1764, 
and not E. tessellatus Linnaeus, 1758. (Malloea is the name of a 
town in Thessaly and is feminine.) 

Dido, new genus. 

TypEe: Dido macrocera (Castelnau) (= Tetralobus macrocerus 
Castelnau, Revue Entomologique, vol. 4, p. 12, 1836). (Here 
designated.) 

Nore: This genus includes only the genotype, a species found in 
Madagascar, and belongs to the subtribe Dicrepidiina. Both Candéze 
(Catalogue Méthodique des Elatérides, p. 49, 1891) and Schenkling 
(Coleopterorum Catalogus, pt. 88, p. 74, 1927) used the generic name 
Coresus for T. macrocerus Castelnau. However, Gemminger and 
Harold (Catalogus Coleopterorum, vol. 5, p. 1509, 1869) validated 
the Dejean name Coresus (MS) as a synonym of Piezophyllus Hope, 


ELATERID BEETLES OF THE WORLD—ARNETT 601 


1842. The genotype of Piezophyllus is P. spencei Hope, designated 
by Hyslop (Proc. U. S. Nat. Mus., vol. 58, pp. 621-680, 1921), and 
it is, therefore, automatically the genotype of Coresus. P. spencei is 
not congeneric with 7. macrocerus, making this new generic name 
necessary. The characters of this genus are as given in Schwarz 
(Genera Insectorum, fasc. 46, p. 61, 1906) under the name Coresus. 
(Dido is a classical Latin name from Greek and is feminine.) 
Mesembria, new genus. 

Tyrer: Mesembria_ subtilis (Candéze) (=Cosmesus subtilis 

Candéze), Monographie des Elatérides, vol. 4, p. 355, 1863. 
(Here designated.) 

Nors: This genus is represented by 62 species, all found in South 
America and all placed in the genus Cosmesus by Schwarz (Genera 
Insectorum, fasc. 46, p. 265, 1907) and Schenkling (Coleopterorum 
Catalogus, pt. 88, pp. 438, 439, 1927). The genotype of Cosmesus 
Eschscholtz, 1829, is C. bilineatus Eschscholtz, 1829, and is not con- 
generic with the species included in Mesembria. Cosmesus Eschscholtz 
is the prior name for Parapomachilius Schwarz, 1900. (Mesembria 
is a classical Latin name and is feminine.) 


Supplement and corrections 


Abelater Fleutiaux 
1947. Notes d’Ent. Chinoise, vol. 11, p. 379. 
Type: Melanozanthus rubiginosus Candéze, Elatérides Nouveaux, 
pt. 2, p. 29, 1878. (Present designation.) 
Invalid genus, no genotype cited; here validated, assuming no previous 
designation. 
Abiphis Fleutiaux 
1926. Bull. Soc. Ent. France, p. 91. 
Type: Hlater nobilis Uliger, ar» Wiedemann, Arch. Zool., vol. 1, 
p. 116, 1800. (Original designation.) 
Abseus Fleutiaux 
1931. Bull. Mus. Hist. Nat., Paris, ser. 2, vol. 3, pp. 611, 
613. 
Type: Anchastus mazimus Fleutiaux, ibid., p. 613. (Original 
designation.) 
Acanthus Van Dyke (not Block, 1795, Borkhausen, 1797, Dumont, 
1816, Gistel, 1834, or Lockington, 1877) 
1932. Proc. California Acad. Sci., ser. 4, vol. 20, p. 387. 
In error for Athous Eschscholtz, 1829. 
Adelocera Latreille 
1829. Crustacés, Arachnides et Partie des Insectes. Jn Cuvier, 
Le Régne Animal... , ed. 2, vol. 4, p. 451, April 1829. 


602 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Type: Elater ovalis Germar, Insectorum Species . . . Vol. 1, 
Coleoptera, p. 49, 1824. (Designated by Hyslop, 1921.) 
This genus is distinct from Agrypnus Eschscholtz, 1829 (Jan.). 
Aeleoderma Fleutiaux 
1932. Mem. Mus. Zool. Univ. Coimbra, ser. 1, No. 55, p. 15. 
In error for Aeoloderma Fleutiaux, 1928. 
Aelus Chagnon 
1935. Nat. Canadien, vol. 62, p. 138. 
In error for Aeolus Eschscholtz, 1829. 
Aemidioides Fleutiaux 
1922. Bull. Mus. Hist. Nat., Paris, vol. 28, p. 47. 
Typn: Aemidioides rohan-chaboti Fleutiaux, loc. cit. (Mono- 
basic.) 
Aeoloderma Fleutiaux 
1928. Encyclopédie Entomologique, ser. B, pt. 1, vol. 3, p. 135. 
Type: Elater crucifer Rossi, Fauna Etrusca, vol. 1, p. 183, 1790. 
(Original designation.) 
Agriodratus Reitter 
1911. Fauna Germanica, vol. 3, p. 222. 
Typsr: Elater pallidulus Mliger, Mag. Insekt., vol. 6, p. 6, 1807. 
(Original designation.) 
Agriotelia Brown 
1933. Canadian Ent., vol. 65, p. 179. 
Typr: Elater bigeminatus Randell, Boston Journ. Nat. Hist., 
vol. 2, p. 37, 1838. (Original designation.) 
Agrypnus Eschscholtz 
1829. Jn Thon, Ent. Archiv, vol. 2, No. 1, p. 32, January 1829. 
Types: Elater murinus Linnaeus, Systema Naturae, ed. 10, p. 
406, 1758. (Designated by Westwood, Introduction to the 
Modern Classification of Insects, vol. 2, synopsis, p. 26, 1838.) 
Aiolus Wilson 
1941. Florida Ent., vol. 24, p. 26. 
In error for Aeolus Eschscholtz, 1829. 
Alestrus Méquignon 
1942. Bull. Soc. Ent. France, vol. 47, p. 10. 
Type: Elastrus dolosus Crotch, Proc. Zool. Soc. London (1867), 
p. 386, 1867. (Monobasic and original designation.) 
Anchastelater Fleutiaux 
1928. Encyclopédie Entomologique, ser. B., pt. 1, vol. 3, p. 144. 
Typr: Anchastelater ornatus Fleutiaux, loc. cit. (Original desig- 
nation and monobasic.) 
Anischia Fleutiaux 
1896. Mém. Soe. Zool. France, p. 300. 


ELATERID BEETLES OF THE WORLD—ARNETT 603 


Type: Anischia boliiana Fleutiaux, loc. cit. (Designated by 
Fleutiaux, Ann. Soc. Ent. France, vol. 105, p. 292, 1936.) 
Fleutiaux designated Anischia languarioides Fleutiaux, 1931, as 
genotype of Anischia (Bull. Ann. Soc. Ent. Belgique, vol. 71, p. 
80, 1931) but the designation is invalid because A. languarioides 
was not an originally included species. 
Antoligostethus Blackburn 
1911. Trans. Roy. Soc. South Australia, vol. 35, p. 203. 
Typs: Antoligostethus lucidus Blackburn, loc. cit. (Monobasic.) 
Aphanopenthes Fleutiaux 

1932. Ann. Soc. Ent. France, vol. 101, p. 35. 

Type: Aphanopenthes acutipennis (Germar) (=Aphanobius 
acutipennis Germar), Zeitschr. Ent., vol. 5, p. 187, 1844. 
(Present designation.) 

Ascoliocerus Méquignon 

1930. Bull. Soc. Ent. France, p. 94. 

Type: Hypnoidus basalis Motschulsky, Mél. Biol. Acad. St. 
Petersbourg, vol. 3, p. 227, 1859. (Original designation.) 

This is a new name for the subgenus Scoliocerus Motschulsky, 1859, 
not Wollaston, 1854. 
Athoina Reitter 

1906. In Heyden, Reitter, and Weise, Catalogus Coleopterorum 
EKuropae ... , ed. 2, p. 399. 

Type: Athous reveliert Mulsant, Ann. Soc. Linn. Lyon, p. 416, 
1874. (Present designation.) 

Athous Eschscholtz 

1829. Jn Thon, Ent. Archiv, vol. 2, No. 1, p. 33: 

Type: Elater vittatus Fabricius, Entomologia Systematica, vol. 1, 
pt. 2, p. 224, 1792. (Designated by Westwood, Introduction 
to the Modern Classification of Insects, vol. 2, synopsis, p. 26, 
1838.) 

Atoloderma Fleutiaux 

1934. Bull. Soc. Ent. France, vol. 39, p. 182. 
In error for Aeoloderma Fleutiaux, 1928. 
Aulacon Arnett 

1952. Wasmann Journ. Biol., vol. 10, p. 112. 

Type: Lepidotus (Aulacon) nobilis (Fall) (=Adelocera nobilis 
Fall) Canadian Ent., vol. 64, p. 58, 1932. (Original designa- 
tion.) 

Betarmou Schenkling 
1927. Coleopterorum Catalogus, pt. 88, p. 543. 
In error for Betarmon Kiesenwetter, 1863. 


604 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Blaiseus Fleutiaus 
1931. Bull. Soc. Zool. France, vol. 56, pp. 307, 308. 
Type: Blaiseus bedeli Fleutiaux, ibid., p. 308. (Monobasic.) 
Brachygonus Buysson 
1912. Bull. Soc. Roumaine Sci., vol. 21, p. 139. 
Types: Brachygonus megerlei (Boisduval and Lacordaire) (= Elater 
megerlet Boisduval and Lacordaire), Faune Entomologique 
... Paris, p. 656, 1835. (Monobasic.) 
Brevicerus Fleutiaux 
1940. Bull. Ann. Soc. Ent. Belgique, vol .80, p. 102. 
Type: Brevicerus cylindricus Fleutiaux, loc. cit. (Original desig- 
nation.) 
Bruyantius Fleutiaux 
1925. Bull. Soc. Ent. France, p. 101. 
Type: Bruyantius capensis Fleutiaux, ibid., p. 102. (Monobasic.) 
Buffeventius Fleutiaux 
1925. Bull. Soc. Ent. France, p. 102. 
Types: Buffeventius lividus Fleutiaux, ibid., p. 103. (Monobasic.) 
Cadichypnus Miwa 
1930. Trans. Nat. Hist. Soc. Formosa, vol. 20, p. 11. 
In error for Cardiohypnus Fleutiaux, 1928. 
Calostirus Thomson 
1864. Skandinaviens Coleoptera, vol. 6, p. 67. 
In error for or isogenotypic with Anostirus Thomson, 1859, p. 103. 
Campsosdernus Fleutiaux 
1927. Faune des Colonies Frangaises, vol. 1, pt. 3, p. 116. 
In error for Campsosternus Latreille, 1834. 
Campsosternss Miwa 
1929. Trans. Nat. Hist. Soc. Formosa, vol. 19, p. 241. 
In error for Campsosternus Latreille, 1834. 
Candezella Szombathy 
1910. Ann. Mus. Nat. Hungarici, vol. 8, p. 354. 
Type: Candezella horvathi Szombathy, ibid., p. 356, fig. 1. 
(Present designation.) 
Cardiohpynus Fleutiaux 
1929. Bull. Soc. Ent. France, p. 23. 
In error for Cardiohypnus Fleutiaux, 1928. 
Cardiohypnus Fleutiaux 
1928. Bull. Soc. Ent. France, p. 284. x 
Tyre: Cardiophorus mirabilis Candéze, Monographie des Ela- 
térides, vol. 3, p. 142, 1860. (Original designation.) 


ELATERID BEETLES OF THE WORLD—ARNETT 605 


Cardiophorus Eschscholtz 

1829. Jn Thon, Ent. Archiv., vol. 2, No. 1, p. 34. 

Tyre: EHlater thoracicus Fabricius, Systema Entomologiae, p. 214, 
1775. (Designated by Westwood, Introduction to the Modern 
Classification of Insects, vol. 2, synopsis, p. 26, 1838.) 

Cardiotarsus Lacordaire 

1857. Histoire Naturelle des Insectes. Genera des Coléoptéres, 
vol. 4, pp. 168, 192. 

Type: Cardiotarsus capensis Lacordaire, ibid., p. 193. (Mono- 
basic.) 

Catelanus Fleutiaux 

1942. Ann. Soc. Ent. France, vol. 111, p. 112. 

Tyre: Hemirhipus trilineatus Castelnau, Revue Entomologique, 
vol. 4, p. 12, 1836. (Monobasic.) 

Cavicoxum Pic 

1928. Mél. Exot.-Ent., vol. 51, p. 21. 

Type: Cavicoxum monstrosum Pic, loc. cit. (Monobasic.) 
Ceroleptus Fleutiaux 

1927. Faune des Colonies Frangaises, vol. 1, pt. 3, pp. 121, 122. 

Typxz: Ceroleptus brevicollis (Candéze), Elatérides Nouveaux, pt. 
2, p. 10, 1878. (Present designation. ) 

Ceropectus Fleutiaux 

1927. Faune des Colonies Frangaises, vol. 1, pt. 3, pp. 110, 117, 
118. 

Typu: Pectocera messi Candéze, Révision de la Monographie des 
Elatérides, fase. 1, p. 207, 1874. (Monobasic.) 

Chalcolepidinus Pjatakowa 

1941. Deutsche Ent. Zeitschr., p. 104. (Publication not seen.) 
In error for Chalcolepidius Eschscholtz, 1829? 
Chatanayus Fleutiaux 

1940. Ann. Soc. Ent. France, vol. 108, pp. 122, 124. 

TypE: Agonischius ruficollis Fleutiaux, Ann. Soc. Ent. France, 
vol. 87, p. 267, 1918. (Monobasic.) 

Chesotraxus Fleutiaux 

1940. Bull. Ann. Soc. Ent. Belgique, vol. 80. pp. 98, 99. 

Typr: Chesotraxus celebensis Fleutiaux, ibid., pp. 99,100. (Orig- 
inal designation. ) 

Chiagosnius Fleutiaux 

1940. Ann. Soc. Ent. France, vol. 108, p. 136. 

Type: Elater obscuripes Gyllenhal. Jn Schoenherr, Synonymia 
Insectorum ... , vol. 3, append., p. 131, 1817. (Original 
designation. ) 


606 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Cleniocerus Stephens 

1829. The Nomenclature of British Insects, p. 10. 

Type: Hlater pectinicornis Linnaeus, Systema Naturae, ed. 10, 
p. 405, 1758. (Designated by Lane, Proc. Ent. Soc. Wash- 
ington, vol. 50, p. 182, 1948.) 

Colaulon Arnett 

1952. Wasmann Journ. Biol., vol. 10, p. 116. 

TypsE: Colaulon rectangularis (Say) (= Elater rectangularis (Say)), 
Ann. Lyc. Nat. Hist. New York, vol. 1, p. 263, 1825. (Original 
designation.) 

Conobajulus Van Zwaluwenberg 

1940. Occ. Pap. Bishop Mus., vol. 16, p. 95. 

Typx: Conobajulus ugiensis Van Zwaluwenberg, ibid., p. 96. 
(Original designation and monobasic.) 

Coresus Gemminger and Harold 

1869. Coleopterorum Catalogus, vol. 5, p. 1509. 

Typr: Piezophyllus spencer Hope, Proc. Zool. Soc. London (1842), 
p. 76, 1842. (Validation in synonymy of Piezophyllus, takes 
same genotype as Prezophyllus.) 

Coryleus Fleutiaux 

1942. Ann. Soc. Ent. France, vol. 111, p. 94. 

Type: Coryleus desruisseauxi Fleutiaux, loc. cit. (Monobasic.) 
Cratonychus Dejean 

1833. Catalogue de la Collection de Coléoptéres .. ., ed. 3, 


p. 87. 
Typr: Elater obscurus Olivier, Entomologie, ou Histoire Naturelle 
des Insectes . . . , Coléoptéres, vol. 2, No. 31, p. 29, pl. 8, 


fig. 6, 1790. (Designated by Blanchard, Histoire des Insectes, 
vol. 2, p. 76, 1845.) 
Crypnoidus Fleutiaux 
1928. Bull. Soc. Ent. France, p. 252. 
TypE: Quasimus setosus Buysson, Coléopt. Rundschau, vol. 3, 
p. 42, 1914. (Original designation.) 
Cryptohypnus Eschscholtz 
1830. Quatember, vol. 2, No. 3, p. 17. 
Type: EHlater riparius Fabricius, Entomologia Systematica, vol. 1, 
No. 2, p. 232, 1792. (Designated by Thomson, Skandinaviens 
Coleoptera, vol. 1, p. 106, 1859.) 
Cryptypnus Munster 
1935. Norsk Ent. Tidsskr., vol. 3, pp. 362-369. (Kmendation 
for Cryptohypnus Eschscholtz, 1833.) 


ELATERID BEETLES OF THE WORLD—ARNETT 607 


Csikia Szombathy 

1910. Ann. Mus. Nat. Hungarici, vol. 8, p. 359. 

TypxE: Csikia dimatoides Szombathy, ibid., p. 360. (Monobasic.) 
Ctenicera Latreille 

1829. Crustacés, Arachnides et Partie des Insectes. Jn Cuvier, 

Le Régne Animal .. . , ed. 2, vol. 4, p. 454, April 1829. 

Type: Hlater pectinicornis Linnaeus, Systema Naturae, ed. 10, 

p. 406, 1758. (Designated by Lane, Proc. Ent. Soc. Wash- 
ington, vol. 50, p. 182, 1948.) 
Ctenicerus Stephens 
1830. Illustrations of British Entomology . . . Mandibulata, 
vol. 3, p. 264. 
Type: Hlater pectinicornis Linnaeus, Systema Naturae, ed. 10, 
p. 405, 1758. (Designated by Westwood, Introduction to the 
Modern Classification of Insects, vol. 2, synopsis, p. 26, 1838.) 
Cteniocerus Miwa 
1934. Rep. Dep. Agric. Gov. Res. Inst. Formosa, No. 65, p. 167. 
In error for Ctenicera Latreille, 1829? 
Ctenocera Stepanov 
1935. Ent. Nachricten, vol. 9, pp. 187-189. 
In error for Ctenicera Latreille, 1829. 
Ctenonyphus Castelnau 
1836. Revue Entomologique, vol. 4, p. 7. 
In error for Ctenonychus Stephens, 1830 (ibid., p. 6). 
Curtisius Miwa 
1934. Rep. Dep. Agric. Gov. Res. Inst. Formosa, No. 65, 
pp. 23, 24. 
Tyres: Aplotarsus maritimus Curtis, Ann. Mag. Nat. Hist., vol. 5, 
p. 277, 1840. (Original designation.) 
New name for Aplotarsus Curtis, 1854, not Stephens, 1830. 
Dactylosimus Fleutiaux 

1922. Trans. Ent. Soc. London, pp. 429, 430. 

Tyee: Dactylosimus dorsalis Fleutiaux, ibid., p. 431. (Monobasic.) 
Dalopius Eschscholtz 

1829. Jn Thon, Ent. Archiv, vol. 2, No. 1, p. 34. 

Typs: Elater marginatus Fabricius, Systema Eleutheratorum 
..., Vol. 2, p. 236, 1801. (Designated by Westwood, 1838.) 
Delox Quelle 

1932. Coleopt. Centralblatt, vol. 5, p. 208. 

TypxE: Adelocera conspersa (Gyllenhall) (=Elater conspersa Gyl- 

lenhall), Insecta Suecica, vol. 1, p.377. (Present designation.) 
Isogenotypic with Danosoma Thomson, 1864. 


809150—55--—2 


608 _. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Diacanthus Latreille (not Rudolphi, 1819) 

1834. Ann. Soc. Ent. France, ser. 1, vol. 3, p. 151. 

Typr: Elater aeneus Linnaeus, Systema Naturae, ed. 10, p. 406, 
1758. (Designated by Thomson, Skandinaviens Coleoptera. 
vol. 1, p. 102, 1859.) 

Isogenotypic with Aphotistus Kirby, 1837. 
Dichonychus Méquignon 
1931. Bull. Soc. Ent. France, p. 207. 
In error for Dicronychus Castelnau, 1840. 
Dicronychus Brullé (not Castelnau, 1840) 
1832. Arachnides—Myriapodes—Insectes—Annélides, in Expédi- 
tion Scientifique de Morée, vol. 3, pt. 1, sec. 2, p. 138. 

Type: Hlater obesus Brullé, loc. cit. (Present designation.) 

This genus is distinct from Dicronychus Castelnau, 1840, which is a 
synonym of Lanelater Arnett, 1952. 
Diocarphus Fleutiaux 

1947. Notes d’Ent. Chinoise, vol. 11, p. 364. 

Type: Phorocardius solitarius Fleutiaux, Bull. Soc. Zool. France, 
vol. 56, pp. 309, 310, 1931. (Monobasic.) 

Diphyaulon Arnett 

1952. Wasmann Journ. Biol., vol. 10, p. 111. 

Type: Lepidotus (Diphyaulon) pyrsolepis (LeConte) (=Adelocera 
pyrsolepis LeConte) Proc. Acad. Nat. Sci. Philadelphia, vol. 
18, p. 389, 1866. (Original designation.) 

Diplocenus Fleutiaux 
1933. Ann. Soc. Ent. France, vol. 102, p. 208. 
In error for Diploconus Candéze, 1860. 
Discrepeidius Van Dyke 
1932. Proc. California Acad. Sci., ser. 4, vol. 20, p. 325. 
In error for Dicrepidius Eschscholtz, 1829. 
Dolopus Méquignon 
1930. Bull. Soc. Ent. France, p. 94. 
In error for Dalopius Eschscholtz, 1829. 
Domenephus Fleutiaux 

1932. Bull. Mus. Hist. Nat., Paris, ser. 2, vol. 4, p. 277. 

Tyre: Phedomenus flavangulus Candéze, Ann. Soc. Ent. Belgique, 
vol. 39, p. 1895. (Designated by Fleutiaux, Journ. East 
Africa Uganda Nat. Hist. Soc., vol. 12, p. 104, 1935.) 

Dresterius Brullé 

1832. Arachnides—Myriapodes—Insectes—Annélides, im HExpédi- 

tion Scientifique de Morée, vol. 3, pt. 1, sec. 2, p. 141. 
In error for Drasterius Eschscholtz, 1829. 


ELATERID BEETLES OF THE WORLD—ARNETT 609 


Endicronychus Fleutiaux 

' 1935. Journ. East Africa Uganda Nat. Hist. Soc., vol. 12, p. 95. 
In error for Hudicronychus Méquignon, 1931. 
Eudicronychus Méquignon 

1931. Bull. Soc. Ent. France, p. 207. 

Type: Dicronychus serraticornis Castelnau, Histoire Naturelle 
des Insectes, vol. 1, Coléoptéres, p. 251, 1840. (Original 
designation.) 

Eupsephus Fleutiaux 

1935. Coleoptera V. Elateridae, in Mission Scientifique de 
V’Omo, vol. 2. Mém. Mus. Hist: Nat., Paris, new ser., vol. '2, 
p. 204. Parts of Mission Scientifique de l’Omo were issued as a 
separate series. 

Typz: Hupsephus dilaticollis Fleutiaux, loc. cit. (Original desig- 
nation.) 

Exophthalmus Berthold (not Schoenherr, 1823) 

1827. Latreille’s Nattirliche Familien des Thierreichs, p. 335. 

Tyre: Hlater linearis Linnaeus, Systema Naturae, ed. 10, p. 404, 
1758. (Designated by Hyslop, Proc. U.S. Nat. Mus., vol. 58, 
p. 646, 1921, as Exophthalmus Latreille, 1829, which is a later 
reference to this same genus. ) 

Fleutiauxellus Méquignon 

1930. Bull. Soc. Ent. France, p. 95. 

Type: Hypnoidus maritimus Curtis, Ann. Nat. Hist., vol. 5, 
p. 277, 1840. (Original designation.) 

Fusimorphus Fleutiaux 

1942. Ann. Soc. Ent. France, vol. 111, p. 111. 

Type: Hemirhipus submetallicus Fleutiaux, Bull. Soc. Ent. 
France, p. 184, 1924. (Monobasic.) 

Gamepenthes Fleutiaux 

1928. Encyclopédie Entomologique, ser. B, pt. 1, vol. 3, p. 158. 

Typm: Megapenthes octomaculatus Schwarz, Deutsche Ent. 
Zeitschr., p. 144, 1898. (Original designation.) 

Ganoxanthus Fleutiaux 

1928. Encyclopédie Entomologique, ser. B., pt. 1, vol. 3, p. 156. 

Typx: Melanoxanthus virgatus Candéze, Monographie des Elaté- 
rides, vol. 2, p. 516, 1859. (Original designation.) 

Georgicus Gistel 
1948. Naturgeschichte des Thierreichs. . . , p. 190. 
Typxn: Georgicus sanguinipennis Gistel, loc. cit. (Monobasic.) 


610 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 108 


Geraniella Gourley 
1950. Trans Proc. Roy. Soc. New Zealand, vol. 78, p. 192. 
(New name for Geranus Sharp, 1877, not Bonaparte, 1854.) 

Gnathodicrus Fleutiaux 

1934. Bull. Soc. Ent. France, vol. 39, pp. 183, 184. 

Tyre: Gnathodicrus francki Fleutiaux, loc. cit. (Monobasic.) 
Gonodyrus Fleutiaux 

1923. Trans. Ent. Soc. London (1922), p. 429. 

Typs: Gonodyrus tarsalis Fleutiaux, ibid., p. 430. (Monobasic.) 
Gyrus Jacobson 

1913. Zhuiki Rossicae, fasc. 10, p. 740. (Publication not seen.) 

Typs: Selatosomus guitatus (Germar) (=Diacanthus guttatus 
Germar), Fauna Insectorum Europae, vol. 21, No. 5, 1817. 
(Present designation.) 

Haupathesus Miwa 
1928. Insecta Matsumurana, vol. 2, p. 133. 
In error for Hapatesus Candéze, 1863. 
Hemirhaphes Fleutiaux 
1930. Bull. Mus. Hist. Nat., Paris, ser. 2, vol. 2, p. 639. 
In error for Hemirrhaphes Candéze, 1878. 
Hemirhipus Berthold 

1827. Latreille’s Natiirliche Familien des Thierreichs, p. 336. 

Type: Elater lineatus Olivier, Entomologie, ou Histoire Naturelle 
des Insectes . . . , Coléoptéres, vol. 2, No. 31, p. 10, pl. 6, fig. 
63, 1790. (Monobasic.) 

Hoabinh Fleutiaux 

1940. Bull. Soc. Zool. France, vol. 65, p. 195. 

Typr: Hoabinh coomani Fleutiaux, loc. cit. (Monobasic.) 
Horizoteichos Van Zwaluwenburg 

1931. Proc. Hawaii Ent. Soc., vol. 7, p. 482, 433. 

Typz: Horizoteichos papuensis Van Zwaluwenburg, ibid., p. 432. 
(Monobasic and original designation.) 

Hypdonus Fleutiaux 

1928. Bull. Soc. Ent. France, p. 149. 

Tres: Hypnoidus bakeri Fleutiaux, Philippine Journ. Sci., ser. 
p., vol. 9, p. 446, 1914. (Monobasic.) 

Hypnoidus Dillwyn 

1829. Memoranda Relating to Coleopterous Insects in the 
Neighborhood of Swansea, p. 32. 

Typ: Elater riparius Fabricius, Entomologia Systematica, p. 
232, 1792. (Designated by Westwood, Introduction to the 
Modern Classification of Insects, vol. 2, synopsis, p. 26, 1838.) 

Hypolittus Weber 
1950. Trans. American Ent. Soc., vol. 76, p. 183. 
In error for Hypolithus Eschscholtz, 1829. 


ELATERID BEETLES: OF THE WORLD—ARNETT 611 


Idiotarmon Binaghi 

1940. Bull. Soc. Ent. Italiana, vol. 72, p. 101. 

Typse: Betarmon quadrivitiatus Ragusa, Nat. Siciliano, vol. 12, 
p. 305, 1893. (Original designation and monobasic.) 

Ipostirus Binaghi 

1940. Mem. Soc. Ent. Italiana, vol. 19, p. 211. 

Tyre: Ipostirus parumcostatus (Buysson) (=Ludius parum- 
costatus Buysson), Faune Gallo-rhénane . . ., vol. 5, p. 80, 
1894. (Original designation.) 

Jonthadocerus Buysson 
1918. Bull. Soc. Hist. Nat. Afrique Nord, vol. 9, p. 109. 
Typr: Jonthadocerus theryi Buysson, ibid., p. 110. (Monobasic.) 
Kudius Miwa 
1928. Insecta Matsumurana, vol. 2, p. 140. 
In error for Ludius Berthold? 
Lacais Fleutiaux 

1942. Ann. Soc. Ent. France, vol. 111, p. 109. 

Typs: Lacais glauca (Castelnau) (=Iphis glauca Castelnau), 
Rev. Ent., vol. 4, p. 9, 1836. (Present designation.) 

Possibly should be considered monobasic, but not clear. 
Lamononia Van Zwaluwenburg 

1928. Insects of Samoa, vol. 4, pt. 2, p. 118. 

Tyre: Lamonona monticola Van Zwaluwenburg, loc. cit. (Mono- 
basic.) 

Lampropsephus Fleutiaux 

1928. Encyclopédie Entomologique, ser. B., pt. 1, vol. 3, p. 104. 

Tyre: Psephus cyaneus Candéze, Elatérides Nouveaux, vol. 2, 
p. 11, 1878. (Monobasic.) 

Lanelater Arnett 

1952. Wasmann Journ. Biol., vol. 10, p. 105. 

Type: Lanelater schotti (ueConte) (=Agryphus schotti LeConte), 
Trans. American Philos. Soc., vol. 10, p. 492, 1853. (Original 
designation.) 

Lasiocerus Buysson 

1912. Bull. Soc. Ent. France, p. 129. 

Types: Cardiophorus schusteri Buysson, ibid., p. 128. (Present 
designation.) 

Lepidotus Stephens (not Asso, 1801, Anales Cien. Nat. (Madrid)) 

1830. Illustrations of British Entomology . . . Mandibulata, 
vol. 3, p. 374. 

Zalepia Arnett, 1953, replaces this homonym. 


612 ~ PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 168 


Lesnelater Fleutiaux 
1935. Journ. East Africa Uganda Nat. Hist. Soc., vol. 12, p. 116. 
Type: Pachyelater madagascariensis Lesne, Bull. Soc. Ent. 
France, p. 173, 1906. (Original designation.) 
New name for Pachyelater Lesne, 1906 (not Lesne, 1897) ibid., p. 174. 
Lincydrus Fleutiaux 
1932. Bull. Soc. Ent. France, vol. 37, p. 148. 
Tyre: Lincydrus cylindricus Fleutiaux, ibid., p. 149. (Mono- 
basic.) 


Lobitarsus Fleutiaux 
1935. Journ. East Africa Uganda Nat. Hist. Soc., vol. 12, p. 93. 
In error for Lobotarsus Schwarz, 1898. 
Ludius Berthold 
1827. Latreille’s Natiirliche Familien des Thierreichs, p. 336. 
Type: Elater ferrugineus Linnaeus, Systema Naturae, ed. 10, 
_p. 405, 1758. (Monobasic.) 
Magacnemis Hyslop 
1921. Proc. U.S. Nat. Mus., vol. 58, p. 655. 
In error for Megacnemis Schwarz, 1906. 
Malekula Van Zwaluwenburg 

1940. Occ. Pap. Bishop Mus., vol. 16, p. 129. 

Type: Malekula piceus Van Zwaluwenburg, ibid., p. 130. (Origi- 
nal designation and monobasic.) 

Mallerius Fleutiaux 

1933. Bull. Soc. Ent. France, vol. 38, p. 280. 

Tyre: Mallerius brasiliensis Fleutiaux, loc. cit. (Monobasic.) 
Melanatractus Fleutiaux 

1933. Ann. Soc. Ent. France, vol. 102, pp. 200, 201. 

Tyre: Melanatractus acutus Fleutiaux, loc. cit. (Monobasic and 
original designation.) 

Melanotus Eschscholtz 

1829. Jn Thon, Ent. Archiv., vol. 2, p. 32. 

Tyre: Elater fulvipes Herbst, Natursystem .. . Insekten, Kafer, 
vol. 10, p. 46, pl. 162, fig. 2, 1806. (Designated by Westwood, 
Introduction to the Modern Classification of Insects, vol. 2, 
synopsis, p. 26, 1838.) 

Melanoxocanthus Miwa 

1930. Wiener Ent. Zeit., vol. 47, p. 97. 
In error for Melanozanthus Eschscholtz, 1836. 
Meristhus Candéze 

1857. Monographie des Elatérides, vol. 1, p. 162. 

Tyre: Meristhus scobinula Candéze, ibid., p. 164. (Designated 
by Candéze, Elatérides Nouveaux, fasc. 6, p. 12, 1896.) 


ELATERID BEETLES OF THE WORLD—ARNETT 613 


Metactenicerus Miwa 
1934. Rep. Dep. Agric. Gov. Res. Inst. Formosa, No. 65, p. 37. 
Type: Corymhites gratus Lewis, Ann. Mag. Nat. Hist. ser. 6, 
vol. 13, p. 262, 1894. (Original designation.) 
Mopleonus Fleutiaux 
1935. Journ. East Africa Uganda Nat. Hist. Soc., vol. 12, pp. 
114-115. 
Type: Mopleonus candezei (Fleutiaux) (=Nomopleus candezei 
Fleutiaux), Insectes Coléoptéres, in Alluaud and Jeannel, Voy- 
age de Ch. Alluaud et R. Jeannel en Afrique Orientale (1911- 
1912), vol. 13, p. 108, 1919. (Original designation.) 
Neolacon Miwa 
1929. Trans. Nat. Hist. Soc. Formosa, vol. 19, pp. 225, 234-236. 
Type: Neolacon formosanus Miwa, ibid., pp. 235-236. (Original 
designation.) 
Neonomopleus Schenkling 
1927. Coleopterorum Catalogus, pt. 88, pp. 308, 331. 
Type: Neonomopleus strictus Candéze, Monographie des Elaté- 
rides, vol. 4, p. 498, 1863. (Present designation.) 
New name for Nomopleus Reitter, 1891 (not Candéze, 1891.) 
Nomopleus Candéze 
1891. Catalogue Méthodique des Elatérides, p. 213. 
Type: Pleonomus argentatus Candéze, Monographie des Elaté- 
rides, vol. 4, p. 499, 1863. (Present designation.) 
Nuilarborica Blackburn 
1911. Trans. Roy. Soc. South Australia, vol. 35, p. 202. 
Types: Nullaborica concinna Blackburn, loc. cit. (Monobasic.) 
Nyctor Semenov-Tian-Shanskij and Pjatakova 
1936. Ceskoslovensk& Spol. Ent., vol. 33, pp. 101, 102. 
Type: Nyctor expallidus Semenov-Tian-Shanskij and Pjatakova, 
ibid., pp. 102, 103. (Monobasic and original designation.) 
Odontocardus Fleutiaux 
1931. Bull. Soc. Zool. France, vol. 56, pp. 307, 332. 
Type: Cardiopherus vitalisi Fleutiaux, Ann. Soc. Ent. France, 
vol. 87, p. 231, 1918. (Original designation.) 
Ovalpalpus Orellana 
1939. Rev. Chilena Hist. Nat., vol. 42, p. 165. 
In error for Ovipalpus Solier, 1851. 
Pacificola Van Zwaluwenburg 
1932. Bull. Bishop Mus., vol. 98, pp. 131-133. 
Type: Pacificola obscura Van Zwaluwenburg, ibid., pp. 133-135. 
(Original designation.) 


614 - PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Paracapiophonus Fleutiaux 
1929. Bull. Soc. Ent. France, p. 23. 
In error for Paracardiophorus Schwarz, 1895. 
Paradaxon Fleutiaux 
1929. Ann. Soc. Ent. France, vol. 98, p. 224. 
In error for Paradozon Fleutiaux, 1903. 
Paradima Miwa 
1929. Insecta Matsumurana, vol. 3, pp. 36-39. 
Typr: Paradima tattakensis Miwa, op. cit., p. 38. (Monobasic 
and original designation.) 
Parastirus Binaghi 
1940. Mem. Soc. Ent. Italiana, vol. 19, pp. 197, 204. 
Typr: Elater purpureus Poda von Neuhaus, Insecta Musaei 
Graecunsis, p. 41, 1761. (Original designation.) 
Parathous Fleutiaux 
1918. Ann. Soc. Ent. France, vol. 87, p. 242. 
Typr: Parathous sanguineus Fleutiaux, ibid., p. 126. (Mono- 
basic.) 
Paroedostethus Van Dyke 
1932. Proc. California Acad. Sci., ser. 4, vol. 20, p. 327. 
Typr: Paroedostethus relictus Van Dyke, ibid., p. 328. (Mono- 
basic and original designation.) 
Parvistoma Fleutiaux 
1929. Ann. Soc. Ent. France, vol. 98, pp. 230, 233-235. 
Typr: Corymbites tenuicornis Fleutiaux, Bull. Soc. Ent. France, 
p. 138, 1902. (Original designation.) 
Patricia Van Zwaluwenburg 
1947. Proc. Hawaii Ent. Soc., vol. 13, p. 113. 
Typr: Patricia austratica Van Zwaluwenburg, ibid., p. 114. 
(Monobasic and original designation.) 
Pectoerra Miwa 
1929. Trans. Nat. Hist. Soc. Formosa, vol. 19, p. 239. 
In error for Pectocera Hope, 1842. 
Pectora Reymond 
1939. Misc. Ent., ann. 47, vol. 40, p. 34. 
In error for Pectocera Hope, 1842. 
Pengamethes Fleutiaux 
1928. Encyclopédie Entomologique, ser. B., pt. 1, vol. 3, p. 173. 
Type: Pengamethes fulvicollis Fleutiaux, ibid., p. 176. (Original 
designation.) 
Pharotarsus Motschulsky 
1861. Bull. Soc. Nat. Moscou, vol. 34, pt. 1, p. 119. 
In error for Phorotarsus Motschulsky, 1859. 


ELATERID BEETLES OF THE WORLD—ARNETT 615 


Pherhimius Fleutiaux 
1942. Ann. Soc. Ent. France, vol. 111, pp. 112, 114. 
Typsz: Elater fascicularis Fabricius, Mantissa Insectorum .. . , 
p. 171, 1787. (Original designation.) 
Phibisa Fleutiaux 
1942. Ann. Soc. Ent. France, vol. 111, p. 105. 
Type: Ctenicera pupiert Fleutiaux, Bull. Soc. Ent. France, p. 
228, 1903. (Original designation.) 
Phorocardius Fleutiaux 
1931. Bull. Soc. Zool. France, vol. 56, pp. 308-312. 
Type: Cardiophorus florentini Fleutiaux, Ann. Soc. Ent. France, 
vol. 62, p. 687, 1894. (Original designation.) 
Placonides Fleutiaux 
1933. Ann. Soc. Ent. France, vol. 102, p. 235. 
Tn error for Ploconides Fleutiaux, 1933. 
Ploconides Fleutiaux 
1933. Ann. Soc. Ent. France, vol. 102, p. 208. 
Typu: Diploconus spiloderus Candéze, Elatérides Nouveaux, vol. 
1, p. 45, 1857. (Original designation.) 
Practapyrus Fleutiaux 
1929. Ann. Soc. Ent. France, vol. 98, pp. 223, 224, 227, 228, 242. 
Typz: Practapyrus descarpentriesi Fleutiaux, ibid., p. 228. 
(Monobasic and original designation. ) 
Pristolophus Van Dyke 
1932. Proc. California Acad. Sci., ser. 4, vol. 20, p. 442. 
In error for Pristilophus Latreille, 1834. 
Prodrasterius Fleutiaux 
1927. Bull. Soc. Ent. France, p. 91. 
Type: Drasterius brahminus Candéze, Monographie des Elatér- 
ides, vol. 2, p. 426, 1859. (Original designation.) 
Prolacon Fleutiaux 
1934. Bull. Soc. Ent. France, vol. 39, pp. 179, 180. 
Typxr: Prolacon allaudi Fleutiaux, loc. cit. (Monobasic.) 
Proquasimus Fleutiaux 
1932. Soc. Ent. France, Livre de Centenaire, pp. 189, 191. 
TypE: Cryptohypnus micros Fairmaire, Ann. Soc. Ent. France, p. 
204, 1903. (Monobasic.) 
Pseudathous Méquignon 
1930. Bull. Soc. Ent. France, p. 95. 
Typr: Athous hirtus (Herbst) (= Elater hirtus Herbst, Archiv der 
Insectengeschichte, vol. 5, p. 114, 1784). (Original designa- 
tion.) 


616 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Pseudoderomecus Fleutiaux 

1907. Rev. Chiliana Hist. Nat., vol. 11, p. 186. 

Type: Medonia fairmairei Candéze, Elatérides Nouveaux, vol. 2, 
p. 31, 1878. (Monobasic.) 

Pseudo-Elater Heer 
1847. Neue Denkschr. Allgemeine Schweiz. Ges., vol. 8, p. 143. 
(No incl. sp., fossil.) 
Pseudonomopleus Fleutiaux 

1931. Bull. Soc. Ent. France, pp. 29, 30. 

Typr: Pleonomus niger Candéze, Elatérides Nouveaux, vol. 2, 
p. 54, 1878. (Original designation.) 

Pseudopristilophus Méquignon 

1930. Bull. Soc. Ent. France, p. 93. 

Tyre: Pristilophus sericans Germar, Zeitschr. Ent., vol. 4, p. 87, 
1843. (Monobasic and original designation.) 

Pseudostirus Binaghi 

1940. Mem. Soc. Ent. Italiana, vol. 19, pp. 198, 221. 

Type: OCorymbites sulphuripennis Germar, Zeitschr. Ent., vol. 4, 
p. 55, 1843. (Original designation.) 

Pulchronotus Fleutiaux 

1933. Ann. Soc. Ent. France, vol. 102, p. 206. 

Type: Diploconus ornatus Candéze, Ann. Mus. Civ. Stor. Nat. 
Genova, ser. 2, vol. 10, p. 782, 1894. (Original designation.) 

Raphaea Fleutiaux 

1933. Ann. Soc. Ent. France, vol. 102, p. 202. 

Type: Raphaea vitida Fleutiaux, loc. cit. (Monobasic and original 
designation.) 

Rastrocephalus Fleutiaux 

1924. Opusc. Ent. Indochine Frangaise, vol. 2, p. 140. 
In error for Rostricephalus Fleutiaux, 1918. 
Rhaciaspis Arnett 

1952. Wasmann Journ. Biol., vol. 10, p. 121. 

Typ: Elater lepidotus Beauvois, Insectes Recueillis en Afrique 
et en Amérique ..., p. 11, pl. 7, fig. 7, 1805. (Original 
designation.) 

Rismethus Fleutiaux 

1947. Notes d’Ent. Chinoise, vol. 11, pp. 242, 257. 

Typr: Meristhus scabinula (=scobinula) Candéze, Monographie 
des Elatérides, vol. 1, p. 164, 1857. (Original designation.) 

Isogenotypic with Meristhus. 
Roggeveenia Van Zwaluwenburg 

1928. Insects of Samoa, vol. 4, pt. 2, p. 120. 

Type: Roggeveenia buxtoni Van Zwaluwenburg, loc. cit. (Mono- 
basic and original designation.) 


ELATERID BEETLES OF THE WORLD—ARNETT 617 


Rostricephalus Fleutiaux 
1918. Ann. Soc. Ent. France, vol. 87, p. 252. 
Typx: Rostricephalus vitalisi Fleutiaux, ibid., p. 253. (Mono- 
basic.) 
Rygodonus Fleutiaux 
1932. Bull. Mus. Hist. Nat., Paris, ser. 2, vol. 4, pp. 856, 857. 
Type: Dorygonus alluaudi Fleutiaux, ibid., p. 863. (Original 
designation.) 
Rymceobites Fleutiaux 
1936. Ann. Soc. Ent. France, vol. 105, pp. 280, 282. 
Typn: Rymecobites singularis Fleutiaux, ibid., p. 282. (Mono- 
basic and original designation.) 
Semiotinus Pjatakova 
1941. Deutsch. Ent. Zeitschr., p. 107. (Not seen.) 
Sicardius Fleutiaux 
1933. Ann. Soc. Ent. France, vol. 102, p. 201. 
Type: Sicardius longicornis Fleutiaux, loc. cit. (Monobasic and 
original designation. ) 
Sphenicosomus Buysson 
1894. Faune Gallo-rhénane . . . , vol. 5, p. 129. 
In error for Spheniscosomus Schwarz, 1892. 
Sternocampsus Fleutiaux 
1927. Bull. Soc. Ent. France, p. 104. 
Typr: Sternocampsus villosus Fleutiaux, loc. cit. (Monobasic.) 
Subathous Fleutiaux 
1918. Ann. Soc. Ent. France, vol. 86, p. 203. 
Types: Subathous tonkinensis Fleutiaux, loc. cit. (Monobasic.) 
Subathrus Schenkling 
1925. Coleopterorum Catalogus, pt. 80, p. 82. 
In error for Subathous Fleutiaux, 1918. 
Sulcilacon Fleutiaux 
1927. Faune des Colonies Frangaises, pp. 56, 65. 
Typ: Adelocera geographica Candéze, Elatérides Nouveaux, vol. 
1, p. 7, 1865. (Original designation.) 
Sulcimerus Fleutiaux 
1947. Notes d’Ent. Chinoise, vol. 11, p. 255. 

Tyree: Meristhus quadripunctatus Candéze, Monographie des 
Elatérides, vol. 1, p. 163, 1857. (Present designation. ) 
Invalid genus, no genotype cited; here validated, assuming no pre- 

vious designation. 
Taiwanathous Miwa 
1930. Trans. Nat. Hist. Soc. Formosa, vol. 20, pp. 66, 68. 
Typr: Taiwanathous arisanus Miwa, ibid., p. 68. (Monobasic 
and original designation.) 


618 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Tenalomus, Fleutiaux 

1933. Ann. Soc. Ent. France, vol. 102, pp. 215, 234. 

Typs: Tenalomus fulvipennis Fleutiaux, ibid., p. 234. (Mono- 
basic.) 

Thacana Fleutiaux 

1936. Ann. Soc. Ent. France, vol. 105, p. 282. 

Type: Corymbites cambodiensis Fleutiaux, Ann. Soc. Ent. France, 
vol. 87, p. 250, 1918. (Monobasic and original designation.) 

Tinecus Fleutiaux 

1940. Ann. Soc. Ent. France, vol. 108, pp. 122, 124, 125. 

TyprE: Agriotes (Ectinus) gratiosus Fleutiaux, Bull. Soc. Ent. 
France, p. 278, figs. 1, 2, 1925. (Original designation.) 

Trelasus Fleutiaux 
1922. Trans. Ent. Soc. London (1922), p. 420. 
Tyre: Trelasus antennalis Fleutiaux, loc. cit. (Monobasic.) 
Tyloarsus Fleutiaux 
1934. Bull. Soc. Zool. France, vol. 59, pp. 56, 60. 
In error for Tylotarsus Germar, 1840. 
Tyleudacus Fleutiaux 

1923. Trans. Ent. Soc. London (1922), p. 429 (Note). 

Type: Hudactylus wapleri Sallé, Ann. Soc. Ent. France, ser. 3, 
vol. 3, p. 267, 1855. (New name for Hudactylus Sallé (not 
Fitzinger, 1843). (Monobasic.) 

Isogenotypic with Platycrepidius Candéze, 1859. 
Vuilletus Fleutiaux 

1940. Ann. Soc. Ent. France, vol. 108, pp. 122-124. 

Type: Agonischius altus Candéze, Elatérides Nouveaux, pt. 4, 
p. 54, 1889. (Originial designation.) 

Xanthelater Miwa 

1931. Trans. Nat. Hist. Soc. Formosa, vol. 21, p. 259. 

Type: Elater granulipennis Miwa, Trans. Nat. Hist. Soc. For- 
mosa, vol. 19, p. 489, 1929. (Original designation.) 

Xantherater Miwa 

1934. Rep. Dep. Agric. Gov. Res. Inst. Formosa, No. 65, pp. 
48, 197, 198. 

In error for Xanthelater Miwa, 1931. 
Xantholamprus Fleutiaux 

1935. Bull. Ann. Soc. Ent. Belgique, vol. 75, p. 304, August 1935. 

Type: Pantolamprus sulcicollis Schwarz, Deutsche Ent. Zeitschr., 
p. 93, 1896. (Original designation.) 

This genus also proposed as new by Fleutiaux in Journ. East Africa 
Uganda Nat. Hist. Soc., vol. 12, p. 96, March 1936. No genotype 
designated, three included species of which the above is one. 


ELATERID BEETLES OF THE WORLD—ARNETT 619 


Xanthopenthes Fleutiaux 
1928. Encyclopédie Entomologique, ser. B, pt. 1, vol. 3, p. 166. 
Type: Megapenthes birmanicus Candéze, Ann. Mus. Civ. Stor. 
Nat. Genova, p. 617, 1888. (Original designation.) 
Yezodima Miwa 
1928. Insecta Matsumurana, vol. 3, pp. 36, 40. 
Type: Yezodima convexunt Miwa, ibid., pp. 39, 40. (Original 
designation.) 
Zalepia Arnett 
1953. Coleopt. Bull., vol. 7, p. 7. 
New name for Lepidotus Stephens, 1830 (not Asso, 1801). 
Zorochros Thomson 
1859. Skandinaviens Coleoptera, vol. 1, p. 106. 
This is the original spelling, Zorochrus is the emendation. 


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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM 


NSTITUTION 
U. S. NATIONAL MUSEUM 


Vol. 103 Washington: 1955 No. 3337 


NEOTROPICAL MIRIDAE, LXIV: NEW BUGS OF THE 
SUBFAMILY CYLAPINAE (HEMIPTERA) 


By José C. M. Carvarno!' 


Through the courtesy of the U. S. National Museum I was able to 
study specimens of Cylapinae in that institution’s collection, among 
which were found the new genera and species herein described. 


Brachyfulvius, new genus 


Cylapinae, Fulviini. Body rounded, strongly convex on hemielytra, 
distinctly shagreened and very short pubescent. 

Head elongate, protruding between the antennae, somewhat hori- 
zontal, with a very short neck; eyes small, slightly removed from an- 
terior margin of pronotum; vertex sulcate; seen from side, clypeus 
prominent, gula long; rostrum reaching about middle of abdomen, the 
first segment reaching level of base of head, remaining segments de- 
creasing gradually in thickness. 

Antennae inserted in front of eyes, removed from anterior margin 
of the latter by a space equal to thickness of base of first antennal 
segment, the latter thick, incrassate towards the apex, second segment 
linear, about 2.5 times longer than the first, both shortly pilose; third 
and fourth joints very slender and longly pilose. 

Pronotum with a distinct collar, the calli strongly raised and forming 
two conical protuberances with a median furrow, both occupying an 
area of about three-fourths of disc, anterior margin of pronotum curved 
posteriorly, lateral margins rounded and depressed anteriorly, then 


1 Museu Nacional, Rio de Janeiro, Brazil; John Simon Guggenheim Memorial fellow, 1953. Additional 
help granted by the Brazilian National Research Council. 


309151—55 621 


622 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


strongly emarginate, humeral angles prominent, posterior margin 
widely emarginate; mesoscutum broadly exposed; scutellum small with 
a median conical protuberance. 

Hemielytra brachypterous, shagreened, strongly convex at middle, 
without membrane, cuneus absent (in its place there is a calloused area 
over margin of corium), clavus, corium, and embolium fused but 
delineated at base, the embolial area flattened throughout. 

Apex of abdomen, seen from above, showing a short anal tube. Legs 
shagreened, the anterior femora narrowed towards the apex, tibiae 
with short spines and hairs, tarsi long, claws of the Cylapinae type, 
very long and slender. Two last legs mutilated. 

Type of genus, Brachyfulvius chapint, new genus, new species. 

This genus is easily differentiated from others of the tribe Fulviini 
by its brachypterous condition and the strongly raised calli and 
scutellum. At first sight it resembles a small beetle. 


Brachyfulvius chapini, new species 


Ficures 72, 74,a; PLATE 15,a 


Characterized by its color, dimensions, and protuberances of 
pronotum. 

Female: Length 2.8 mm., width 1.5 mm., head length 0.5 mm., 
width 0.6 mm., vertex 0.35 mm. Antennae segment I length 0.3 mm.; 
II, 0.8 mm.; III, 0.8 mm.; IV, 0.4 mm. Pronotum length 0.4 mm., 
width at base 0.9 mm. Rostrum length 1.7 mm.; segment I length 0.42 
mm.; II, 0.45 mm.; JII, 0.42 mm.; IV, 0.48 mm. 

Color dark brownish; head, rostrum, antennae, legs, and embolial 
area lighter; apical callosity of corium and anal tube yellowish; second 
antennal segment whitish at apex; coxae light with a reddish tinge on 
the inner surface. 

Morphological characters as given for genus. Male unknown. 

Holotype: Female, USNM 61939, Cinchona, Jamaica, May 9, 1941, 
K. A. Chapin. 

As in the genus Corcovadocola Carvalho, 1948, whose females are 
brachypterous, the males of Brachyfulvius are expected to be macrop- 
terous. 

I take pleasure in naming this species for Dr. E. A. Chapin, who 
collected this interesting new genus and species of mirid. 


Peritropoides, new genus 


Cylapinae, Fulviini. Species of small size, body ovoid, smooth, 
beset with rigid semierect pubescence. Head inclined and pointed in 
front, vertex convex and inclined towards frons, posterior margin 
straight, frons strongly inclined in the same level of clypeus; eyes 


NEOTROPICAL MIRIDAE, LXIV—CARVALHO 623 


divergent anteriorly, touching anterior angles of pronotum; seen from 
side, the eye is compressed and touches the gula below or nearly so. 
Rostrum reaching middle of abdomen or slightly beyond, the first seg- 
ment thicker than the others, reaching slightly beyond the base of 
anterior coxae, remaining segments about subequal in length. 

Antennae inserted a little above inferior line of orbita, touching 
anterior margin of eye, segment I very short, reaching about apex of 
clypeus, segment II subequal in thickness to the first and three times 
longer, linear, segments III and IV more slender, the fourth twice 
as long as third or so, the whole antenna covered by very short 
pubescence. 


Figure 72.—Brachyfulvius chapini, new genus and new species, female, holotype; 
seen from above. 


Pronotum slightly inclined towards the head, collar present, 
widest at middle and narrowed towards the sides, calli area somewhat 
raised on two anterior thirds of pronotum, lateral margins of disc 
rounded and declivous, posterior margin rounded at lateral angles, 
emarginate in front of mesoscutum; the latter prominent, carinate 
transversally at middle; scutellum relatively small, slightly convex. 


624 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 

Hemielytra smooth, densely pubescent, embolium more distinct 
towards the base, cuneus as long as wide at base, fracture distinct, 
incisure obsolete; membrane biareolate, very minutely pubescent 
(iridescent under reflected light). 

Legs relatively short, the posterior femora incrassate, tibiae with 
slender spines and common pubescence, tarsi very long, claws of the 
Cylapinae type. Cleft of anterior coxae deeply incised. 

Type of genus, Peritropoides annulatus, new species. 


0.5 mri. 


Figure 73.—Fulvius albonotatus, new species, male, holotype. 


This genus is very near Peritropis Uhler, 1891, being easily dis- 


tinguished, however, by the presence of a distinct collar, sides of 
pronotum not carinate, calli not raised, narrow embolium, vertex 
smooth, and body surface not shagreen. 


NEOTROPICAL MIRIDAE, LXIV—CARVALHO 625 


Peritropoides annulatus, new species 


PLATE 15,B 


Characterized by its color, size, and structure of head. 

Female: Length 2.2 mm., width 0.9 mm. Head: Length 0.2 mm., 
width 0.3 mm., vertex 0.22 mm. Antennae: Segment I length 0.1 
mm.; II, 0.4 mm.; ITI, 0.2mm.;IV,0.3mm. Pronotum: Length 0.3 
mm., width at base 0.8 mm. Rostrum: Length 1.0 mm., segment I 
length 0.25 mm.; II, 0.28 mm.; III, 0.25 mm.; IV, 0.25 mm. 

Color brownish with whitish spots or areas; head brown on vertex 
with two whitish spots bordering eye, clypeus and frons with brown 
and whitish areas, sides of head, clypeus, and first segment of rostrum 
reddish brown, remaining rostral segments yellowish; eyes dark 
brown; first antenna brown with whitish apex, second antenna with 
three whitish and two brown rings, third and fourth antennae brown 
with whitish apices; pronotum brown, especially at middle and 
lateral margins, with yellowish spots, the posterior margin with three 
distinct whitish spots, one adjacent to each lateral angle of mesoscutum 
and the third median; mesoscutum also with two whitish spots at base; 
scutellum brown with whitish apex; hemielytra with clavus and basal 
half of cori'um white with small brown dots, apical portion of corium 
and embolium brown to reddish brown (except a white spot on apex of 
embolium and external apical angle of corium reaching the middle of 
base of cuneus), cuneus reddish brown with extreme apex white; 
membrane infumate; underside darkish brown, apex of coxa reddish, 
trochanters and extreme apex of femora whitish, the posterior pair 
noticeably darker, tibiae yellow with a small black ring near base and 
a large one near middle, the anterior pair with a median and two 
basal rings; whitish apex of posterior femur with a reddish to brown 
ring. 

Morphological characters as given for genus. Male unknown. 

Holotype: Female, USNM 61940, Colombia (on orchid), intercepted 
at San Francisco, Calif., Dec. 16, 1940. 


Peritropoides quadrinctiatus, new species 
FicureEs 74,b, 75,e,f, 76,0,f 


Characterized by its color, dimensions, and structure of head. 

Male: Length 3.0 mm., width 1.1 mm. Head: Length 0.3 mm., 
width 0.5 mm., vertex 0.24 mm. Antennae: Segment I length 0.2 
mm.; II, 0.6 mm.; III, 0.8 mm.; 1V, 0.6 mm. Pronotum: Length 
0.4 mm., width at base 1.0mm. Rostrum: Length 1.7 mm.;segment 
I length 0.42 mm.; II, 0.45 mm.; III, 0.42 mm.; IV, 0.45 mm. 


626 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


Color yellowish mottled with brown, reddish marmorate on em- 
bolium, the hemielytra with four brown areas; head yellowish on 
vertex and frons (except for a small central brown spot at middle of 
vertex and two other bordering eye near base of frons), sides of head 
above antennae with a reddish brown fascia reaching from eye to 
clypeus, the latter variegated with red becoming solid reddish brown 


0.5mm, 


Figure 74.—a, Brachyfulvius chapint, new species, side view of head and pronotum; 
b, Peritropoides quadrinotatus, new species, head and pronotum from above; 
c, Fulvius ornatifrons, new species, head and pronotum of female, holotype; 
d, Vannius oculatus, new species, showing color pattern. 


towards the apex, genae and first rostral segment yellowish brown, 
gula brown, the remaining rostral segments yellowish; eyes brown; 
antennae yellowish, the apex of first joint with a reddish ring; pro- 
notum variegated with brown, two roundish brown spots are to be 
seen over the collar and a brownish fascia behind each eye occupying 
also lateral margin of pronotum (on anterior margin, contiguous to 
eye there are two short whitish fasciae below and above the pronotal 
margin, which is also whitish), pleural region bark brown; mesoscutum 
with brown spots at basal angles and a more pronounced dark brown 


U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 103, PLATE 15 


A, Brachyfulvius chapini, new genus and new species, female, holotype. B, Peritropoides 
annulatus, new species, female, holotype. 


NEOTROPICAL MIRIDAE, LXIV—CARVALHO 627 


median portion separated from angles by a whitish raised spot; 
scutellum brown with faint yellowish spots and whitish apex; hemi- 
elytra yellowish, variegated with brown externally and vice versa 
interiorly, clavus with two large median brown spots, embolium 
marmorated with red, corium with a faint brownish area beyond apex 


Figure 75.—Male genitalia: a, pygophore of Fulvius castaneous, new species, left 
side, lateral view; 6, pygophore of F. quadristillatus (Stal), left side, lateral 
view; c, pygophore of F. albonotatus, new species, left side, lateral view; d, 
aedeagus of F’. castaneous, new species; e, apex of pygophore of Peritropoides 
quadrinotatus, new species, seen from above; f, aedeagus of P. quadrinotatus; g, 
pygophore of Fulvius bisbistillatus (Stal), left side, lateral view. 


of cubital vein, cuneus whitish with reddish apex and a well marked 
dark brown spot on inner basal angles; membrane infumate, the veins 
yellowish; underside of body yellowish at median line, the tergites 
becoming darker towards the upper surface, legs yellowish (coxae and 
trochanters), femora and tibiae mutilated. 


628 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


Head more horizontal than in the preceding species, the eyes larger, 
straight posteriorly, coarsely granulose, second antennal segment 
slightly incrassate towards the apex, body pubescence long and semi- 
erect, eyes reaching the gula below. 

Aedeagus (fig. 75, f) of the Cylapinae type, short and thick, without 
chitinized teeth internally. Left clasper (figs. 75, e, 76, 6, f) curved, 
pointed apically, with a median short prong. Right clasper small, 
roundish apically, with a few setae. 

Female unknown. 

Holotype: Male, USNM 61941, Barro Colorado Island, Canal Zone, 
Panamé, July 1923, R. C. Shannon. 

This species differs from Peritropoides annulatus, new species, by 
its color and structure of head. Its trivial name is derived from the 
four brown spots to be seen on clavus and internal angles of cuneus. 


Vannius oculatus, new species 
Figure 74,d 


Characterized by its color, structure of eyes, and yellow first anten- 
nal segment. 

Female: Length 2.7 mm., width 0.8 mm. Head: Length 0.3 mm., 
width 0.4 mm., vertex 0.14 mm. Antennae: Segment I length 0.2 
mm.; II, 1.0 mm.; III, 1.3 mm.; IV, 0.4 mm. Pronotum: Length 
0.2 mm., width at base 0.7 mm. Rostrum: Length 0.71 mm.; seg- 
ment I length 0.17 mm.; II, 0.25 mm.; III, 0.14 mm.; IV, 0.14 mm. 

Color yellowish with red areas; eyes, sides of pronotum, median 
portion of scutellum, corium bordering clavus and a small fascia on 
the latter joining the corial fascia, a round spot on apex of corium and 
a smaller one on intermal margin of cuneus, frons and sides of head, 
pleural region, apex of abdomen, and base of same on upper surface, 
red; antennae (except red ring near apex and whitish apical portion), 
rostrum, coxae, and middle circular band of abdomen, yellow. The 
third antennal segment is fuscous at base. 

Head small, sulcate on frons and vertex, eyes very large, elongate, 
narrowed posteriorly, longer than the head as seen from above; seen 
from side, eyes distant from gula by a space about equal to length of 
eye, lorum strongly enlarged, gula very short. Rostrum reaching very 
slightly beyond the hind coxae. 

Pronotum with anterior margin raised and hooded over the vertex 
between the eyes, emarginate behind each eye, lateral margins 
rounded and slightly emarginate at middle, the posterior margin 
biconcave, humeral angles produced; mesoscutum exposed; scutellum 
elongate with acute apex. 


NEOTROPICAL MIRIDAE, LXIV—-CARVALHO 629 


Hemielytra elongate, the margins subparallel, cuneus very long, 
almost three times as long as wide at base, embolium very narrow, 
linear. Legs mutilated. Male unknown. 

Holotype: Female, USNM 61942, Costa Rica (on bananas), inter- 
cepted at New York, May 11, 1936. 

This species is similar to Vannius rubrovittatus Distant, 1893, but 
easily distinguished by the yellow first antennal segment, the eyes 
pointed posteriorly, and different color markings of apex of corium 
and cuneus. 


Fulvius albonotatus, new species 


Figures 73, 75,c, 76,d,f 


Characterized by its small size, whitish spots of head and pronotum, 
and male genitalia, 

Male: Length 2.4 mm., width 1.0mm. Head: Length 0.3 mm., with 
0.4mm., vertex0.21 mm. Antennae: Segment I length 0.2 mm.; II, 
0.5 mm.; III, 0.2 mm.; IV, 0.4mm. Pronotum: Length 0.2 mm., 
width at base 0.7 mm. Rostrum: Length 1.0 mm., segment I length 
0.25 mm.; IT, 0.28 mm.; III, 0.28 mm.; IV, 0.28 mm. 


Figure 76.—Male claspers: a, Fulvius castaneous, new species, left clasper; }, 
Peritropoides quadrinotatus, new species, left clasper, dorsal view; c, Fulvius 
castaneous, new species, right clasper; d, Ff’. albonotatus, new species, left clasper, 
dorsal view; e, Peritropoides quadrinotatus, new species, left clasper, ventral 
view; f, Fulvius albonotatus, new species, left clasper, side view. 


Color brown, distinctly darker on head and pronotum; head dark 
brown with a whitish, short fascia on vertex between the eyes, pro- 
notum also dark brown with a whitish spot touching posterior margin 
on median line; antenna with segment I reddish brown, lighter at apex 
(but not white), segment IT yellowish sulphurescent, whitish at apex 
and lighter at base, third and fourth segments fuscous; mesoscutum 
dark brown at middle, lighter on basal angles; scutellum dark brown 


630 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 108 


with whitish apex; hemielytra chestnut brown with a roundish small 
white area near base of corium on level with middle portion of clavus, 
cuneus whitish on basal half, reddish brown apically; membrane 
infumate; underside with pleural portion dark brownish, abdomen 
reddish brown, coxae white (except extreme base), femora brown with 
reddish apices, tibiae pale, each with a brown to fuscous ring near base, 
tarsi yellow. 

Head strongly pointed, first antennal segment incrassate, narrowed 
toward the base, second linear, about half as thick as the first, remain- 
ing joints very slender. Pronotum slightly biconcave posteriorly. 
Hemielytra with embolium enlarged towards the apex, cuneus about 
as long as wide. Rostrum reaching beyond posterior coxae. 

Aedeagus similar to that of Fulvius castaneous, new species (fig. 
75, d); left clasper as seen in figures 76,d,f. Pygophore seen from 
left lateral aspect as in figure 75,c. 

Female slightly more robust than male but similar in color and 
general appearence. 

Holotype: Male, USNM 61943, Barro Colorado Island, Canal Zone, 
Panama (on Heliconia mariae flowers), June 1940, J. Zetek, No. 4667. 
Allotype: Female, same data. Paratypes: 9 o'o' and 3 99, same 
data. Also 1 o, Panamé (with Hevea and other seeds), intercepted at 
Hoboken, N. J., Sept. 8, 1950. 

This species differs from all others in the genus by its small size, 
color markings, and male genitalia. It is easily distinguished from 
Fulvius lunulatus Uhler, 1891, by its color pattern of the head and 
pronotum. These structures are uniformly dark in the latter species. 


Fulvius castaneous, new species 


Fieure 75,a,d; 76,a,c. 


Characterized by the very faint whitish area on corium, large size, 
and male genitalia. 

Male: Length 3.9 mm., width 1.5 mm. Head: Length 0.4 mm., 
width 0.6 mm., vertex 0.21 mm. Antennae: Segment I length 0.4 
mm.; II, 1.0 mm.; III, 0.4 mm.; IV, 0.7 mm. Pronotum: Length 
0.5 mm., width at base 1.1mm. Rostrum: Length 2.0 mm., segment 
I length 0.50 mm.; IJ, 0.57 mm.; III, 0.57 mm.; IV, 0.42 mm. 

Color blackish brown; head, pronotum, and antennae (except 
whitish apical one-fourth of second segment) unicolorous; mesoscutum 
with two lighter areas; scutellum unicolorous, dark brown; hemielytra 
brown to cinnamom, with a faint whitish area on corium in level with 
middle of clavus, apex of the latter also lighter, cuneus reddish brown 
apically, white on basal third; membrane infumate; underside black- 
brown on anterior half and reddish brown on abdomen; posterior and 


NEOTROPICAL MIRIDAE, LXIV—-CARVALHO 631 


middle coxae white (except on extreme bases), anterior coxae black 
(except on extreme apex), femora and tibiae brownish, the tibiae 
lighter than femora. In some specimens the whitish mark on corium 
is almost absent, giving the insect the aspect of having a totally black 
corium. 

Body fairly large, head strongly produced, antennae quite long, 
calli small and roundish. 

Aedeagus (fig. 75,d) with a chitinized spiculiform projection inter- 
nally and a small field of chitinized teeth. Left clasper (fig. 76,a) 
curved apically, with a median enlargement bearing setae. Pygophore 
seen from left lateral view as in figure 76,c. 

Female identical to male in color and dimensions. 

Holotype: Male, USNM 61944, Tigambato, Michoac4an, México 
(on orchid), intercepted at Laredo, Tex., Oct. 16, 1948. Allotype: 
Female, México, Distrito Federal, México (on Philodendron sp.), 
intercepted at Laredo, Tex., July 29, 1952. Paratypes: 1@ and 19, 
same data as allotype; 1 9, Tamazunchale, San Luis Potosi, México 
(on orchid), intercepted at Laredo, Tex., Dec. 30, 1948; 1 #@, Huicha- 
can, San Luis Potosi, México (on orchid), intercepted at Laredo, Tex., 
Jan. 29, 1938: 1 9, México (on crysanthemum), intercepted at Browns- 
ville, Tex., Sept. 5, 1942; 1 o&, Guatemala (on bananas), intercepted 
at Philadelphia, Pa., Oct. 21, 1935; 1 9, San José, Costa Rica, 
altitude 1,135 meters, P. Biolley (Fulvius quadristillatus Stal, det. 
A. C. Montandon). Paratypes in collections of U. S. National 
Museum and the author. 

This species runs to F. bisbistilatus (Stal) in Poppius’ key (Acta 
Soc. Sci. Fennicae, vol. 37, pt. 4, p. 32, 1909), but differs from the 
latter by the much less marked light area on corium, larger size, and 
male genitalia (fig. 75,g). It also shows similarity with Fulvius 
imbecilis (Say), 1832, but can be distinguished by the color of 
hemielytra, antennae, and male genitalia. From Fulvius quadristil- 
lotus (Stal), 1860, it differs in the color and male genitalia (fig. 75,0). 


Fulvius ornatifrons, new species 
Figure 75,c 


Characterized by its color, size, and very long rostrum. 

Female: Length 2.7 mm., width 1.1mm. Head: Length 0.4 mm., 
width 0.5 mm., vertex 0.30 mm. Antennae: Segment I length 0.3 
mm.; II, 0.8 mm.; III, 0.2 mm.; IV, 0.4mm. Pronotum: Length 
0.4 mm., width at base 1.0 mm. Rostrum: Length 1.7 mm., all seg- 
ments subequal in length. 

Color brown with a reddish tinge; head and calli variegated with 
red or fuscous as seen in figure 75,c, a small whitish fascia on middle of 


632 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


posterior margin of pronotum touching mesoscutum; hemielytra with 
extreme base of corium, extreme apex of clavus, and a faint mark on 
corium in level with median portion of clavus, a semilunal white spot 
on base of cuneus; antennae brown tending to reddish brown, the 
second segment unicolorous; underside reddish brown, middle and 
posterior coxae white (except at extreme bases), anterior coxae reddish 
brown, whitish at base; legs reddish brown, extreme base and apices 
of femora whitish, tibiae lighter than femora tending to yellow; 
rostrum yellow. 

Species of small size, the antennae fairly thick, calli of pronotum 
noticeably raised, rostrum reaching the genital segment (male). 

Male slightly smaller and more reddish than the female and the 
calli not so distinctly marked. The genitalia of the male was lost 
when being manipulated. 

Holotype: Female, USNM 61945, Caracas, Venezuela (on Catileya 
sp.), intercepted at San Francisco, Calif., July 10, 1946. Paratypes: 
1, Caracas, Venezuela (on Cattleya speciosissima), intercepted at 
San Francisco, Calif., May 23, 1946; 1 o&, Venezuela (on orchid), 
intercepted at Washington, D.C., July 14, 1939. Paratypes in col- 
lections of U. S. National Museum and the author. 

This species differs from all others in the genus by the variegated 
markings of head and pronotum as seen in figure 74,¢. 


U, S$, GOVERNMENT PRINTING OFFICE: 19558 


INDEX 


(New genera, species, etc., are printed in italics. 


Page numbers of principal 


entries also in italics) 


Abbott, R. Tucker; 
Atlantic periwinkles Nodzilittor- 
ina, Echininus, and Tectarius, 
449. 

abbreviata, Amblyops, 581 

Abelater, 601 

aberrans, Stilobezzia (Eukraiohelea), 61 

Abiphis, 601 

abranchiata, Leaena, 209, 325, 326, 

1 (fig.) 
Abseus, 601 
Acanthomysis, 576, 582 
davisi, 582 
macropsis, 582 
nephrophthalma, 582 
pseudomacropsis, 582 
sculpta, 582 
sp., 582 

Acanthus, 601 

accidentalis, Colodon, 430 

acridens, Hyracodon, 430 

Acrostoma, 383 

Actenicerus, 600 

Actinicola, 188 

bicolor, 191 
percula, 191 

acuminata, Strabala, 125 (fig.), 127, 

acuminata, Potamilla, 336 

acuta, Pleurocera, 360 

Pleurocera (Oxytrema), 360 

acutidens, Ictops, 403, 430 

acutifrons, Ampharete, 208, 316, 319 

fi 


(fig. 
Amphicteis, 316 
acutipennis, Aphanobius, 603 
Aphanopenthes, 603 
acutus, Melanatractus, 612 
Pleurocerus (Oxytrema), 360 
Adamatornis, 594 
Adelocera, 601 
conspersa, 607 
geographica, 617 
pyrsolepis, 608 
Adjidaumo, 433 
minimus, 430 
Aeleoderma, 602 
Aelus, 602 
Aemidioides, 602 
rohan-chaboti, 602 
eoepeanis, tiene 87, 88, 89, 91 


aeneus, Elater, 608 
aenigmaticum, Rhopalosoma, 19 
Pinchacus, 471 
Tapirus, 477, 479, 481, 485 


365272—56——2 


Review of the| Aeoloderma, 602, 603 


Aeolus, 602 
Aepinacodon, 434 
americanus, 480 
Aeronautes melanoleucus, 537, 538 
saxatilis saxatilis, 537, 538 
aetherodroma, Chaetura_ spinicauda, 
539, 542 
affinis, Apis affinis, 544 
Flabelligera, 307, 289, 297 (fig.) 
Lophogaster, 577 
Lophogaster subglaber, 577 
Merycoidodon, 430 
africana, Eukraiohelea, 61 
Sternaspis scutata, 309 
Stilobezzia (Eukraiohelea) 63 
agitator, Hapithus, 20, 21, 23, 24, 25, 31 
Hapithus agitator, 23 
Agonischius altus, 618 
ruficollis, 605 
Agriodratus, 602 
Agriotella, 602 
Agriotes (Ectinus) gratiosus, 618 
Agryphus schotti, 611 
Agrypnus, 602 
Aiolus, 602 
akallopisos, Amphiprion, 188, 190, 192 
Prochilus, 188, 190 
alaskensis, Dendrocoelopsis, 165, 168, 
177, 178, 181 (fig.) 
alaskensis, Haploscoloplos, 278, 279 
alatum, Chalaraspidum, 577 
albicincta, Streptoprocne zonaris, 548 
albissima, Phagocata, 165, 167 
albitarsis, Ceratopogon, 138 
albofasciata, Wetmorella, 441, 442, 443, 
446, 447 
albonotatus, Fulvius, 624 (fig.), 627 
(fig.), 629 (fig.) 
Alestrus, 602 
alexandri, Autolytus, 207, 242, 244, 245, 
246 


4 
alfa, Bathygenys, 430 
alluaudi, Dorygonus, 617 
Prolacon, 615 
Allohelea, 137 
polita, 138 
alternata, Syllis, 253, 254 
Typosyllis, 253 
Altica, 121, 122, 123, 124 
ferruginea, 121, 132 
scutellaris, 121 
testacea, 122 
altitalonidus, Apternodus, 430 
altus, Agonischius, 618 
alutaceus, Hyas coarctatus, 261 


633 


634 


alveare, Megare, 364 

Pleurocera, 364 
amarula, Thiara, 378 

Thiara (Thiara), 379 
Ambloxus olequaensis, 360 
Amblyops abbreviata, 581 
Ambrysus, 1 

bispinus, 3 (fig.), 4, 6 

buenoi, 4 

cosmius, 6 

dilatus, 6 

funebris, 1 

guttatipennis, 6 

hungerfordi, 6 

mormon, 3, 4 

parviceps, 6 

pudicus, 6 

puncticollis, 6 

signoreti, 6 

thermarum, 1, 3 (fig.) 

Two new naucorid bugs of the 

genus, 1 

woodburyi, 3 
ambulans, Haltica, 122, 123, 131 

Strabala, 125 (fig.), 127, 131 
Amechanus, 101 

ferrugineus, 102 

fossatus, 103 

serratus, 113 
americana, Mysidella, 576, 582 

Sorocelis, 566 
americanus, Aepinacodon, 430 

Bothriodon, 430 

Lophogaster, 577 

Tapir, 479 

Ceres 476, 477, 479, 480, 481, 

4 

Amotrypane, 296 

breviata, 205, 208, 295, 297 (fig.) 
amnigena, Eukraiohelea, 61 
Ampharete, 315 (key) 

acutifrons, 208, 316, 319 (fig.) 

cirrata, 316 

goési, 208, 317 

goési braznikovi, 317 

grubei, 316 

sp. (young), 317 

trilobata, 316 

vega, 208, 315, 319 (fig.) 
Ampharetidae, 208 (table), 212, 314, 


PROCEEDINGS 


315 (key), 319 (figs.) 
Amphicteis acutifrons, 316 
vega, 315 


Amphictenidae, 310 
Amphicynodontinae, 419, 421 (table), 
434 


Amphicyoninae, 420 
Amphiprion, 187, 188 (key), 192 (table), 
196, 200 


akallopisos, 188, 190, 192 

arion, 199 

bicinctus, 190, 192, 194, 198, 199 
bicolor, 191 

bifasciatus, 193 

bifasciatus annamensis, 193 
boholensis, 198 


OF THE NATIONAL MUSEUM 


VOL. 103 


chrysargyrus, 198 
chrysogaster, 189, 192, 193 
chrysopterus, 198 

clarckii, 198 

de bojer, 198 

ephippium, 190, 192, 199, 201 
ephippium chrysopterus, 195 
frenatus, 190, 192, 199, 200 
fusciventer, 193 

intermedius, 194 

japonicus, 198 

laticlavius, 188, 193 
macrostoma, 201 
mauritiensis, 189, 192, 196 
mecullochi, 201 

melanopus, 190, 192, 201 
melanostolus, 198 
melanurus, 194 
monofasciatus, 199 

ocellaris, 194 

papuensis, 199 

percula, 188, 191, 192, 193 
perideraion, 188, 191, 192 
ae 189, 193, 194, 197, 198, 


Review of the Indo-Pacific anemone 
fishes of the genus, with descrip- 
tions of two new species, 187 
rosenbergi, 191 
rubrocinctus, 199 
riippeli, 199 
sebae, 189, 192, 197 
snyderi, 198 
tricinctus, 189, 192, 195, 197 
tricolor, 199 
trifasciatus, 193 
tunicatus, 191 
unimaculatus, 193 
xanthurus, 190, 192, 198 
Amphitrite, 320, 321 (key) 
cincinnata, 327 
cirrata, 209, 319 (fig.), 321 
groenlandica, 209, 321, 322 
infundibulum, 340 
radiata, 321 
robusta, 322 
Ampullacera maculata, 385 
Ampullarina maculata, 385 
Amyadenium, 177, 178 
brementi, 177 
chattoni, 177 
garmieri, 177 
vandeli, 177 
Anagnota, 560 
Anaitides, 236 
groenlandica, 236 
analis, Euchone, 209, 339, 341 (fig.) 
Larra, 32 
Sabella, 339 
Anaplocamus, 363 
Anatomy and relationships of glossy 
cuckoos of the genera Chryso- 
coccyx, Lampromorpha, and 
Chalcites, 585-597 
Anchastelater, 602 
ornatus, 602 


INDEX 


Anchastus maximus, 601 
Anchialina typica, 580 
Anculosa, 362 
dilatata, 363 
andecolus, Apus andecolus, 536 
andicola, Tapirus, 472 
Angitrema, 363 
Angulidae, 401 
angulus, Bolboceras, 95, 117 
Bolborhombus, 99 (fig.), 115 (map), 
117, 119 
angustata, Stenomicra, 560 
Anischia, 602, 603 
boliviana, 603 
languarioides, 603 
poe ener) Amphiprion _ bifasciatus, 


annectens, Leptomeryx, 430 
annulata, Meronychina, 556 
Microperiscelis, 555, 556 
Myodris, 556 
Notiphila, 552 
Periscelis, 552, 543, 556 
annulatus, Peritropoides, 624, 625, 628 
annulipes, Periscelis, 556 
Anodonta, 368 
anomala, Holmesiella, 580 
Anostirus, 604 
anta, Tapir, 477, 479, 481 
Tapirus, 481, 485 
antarctica, Capitella capitata, 298 
Leaena, 325 
Leaena abranchiata, 325, 326 
Maldane sarsi, 304 
antargucys: Trichobranchus  glacialis, 


antea, Tapirus, 476 

antennalis, Ceratolophus, 68 
Ceratopogon, 68 
Hartomyia, 68 
Johannseniella, 68 
Stilobezzia, 59, 60, 61, 68 


Stilobezzia (Stilobezzia), 68, 73 


Anthias bifasciatus, 194 
clarkii, 198 
polymna, 198 
polymnus, 191 

Anthomyza, 557 

Anthomyzidae, 551, 

(key) 

anthonyi, Eurycaelon, 362 

Anthracotherinae, 434 

anticostiensis, Arcteobia, 206, 226 
Eucranta, 226 
Eupolynoé, 225 
Harmothoé, 225 

Antilopidae, 424 

Antimelania, 381, 382 
costula, 382 
episcopalis, 382 
soriniana, 382 

Antinoé, 214, 215 
badia, 216 
sarsi, 205, 206, 215 


553, 4656, 557 


635 


Antinoélla sarsi, 215 
Antoligostethus, 603 
lucidus, 603 
antoni, Litorina, 458 
saul aera 480, 481, 484, 485, 
4 


Aphanobius acutipennis, 603 
Aphanopenthes, 603 
acutipennis, 603 
Aphrodita cirrhosa, 226 
imbricata, 220 
longa, 230 
minuta, 230 
scabra, 217 
aphroditoides, Castalia, 206, 239, 243 
(fig.) 
Nereis, 239 
Psammate, 239 
Apis affinis affinis, 544 
Aplotarsus, 607 
maritimus, 607 
Apternodus, 431, 433 
altitalonidus, 430 
mediaevus, 403, 430 
Apus andecolus andecolus, 536 
aquaticus, Scalopus, 404 
Archaeolagus sp., 416 
Archaeomysis grebnitzkii, 580 
Archaeotherium, 433 
crassum, 430 
marshi, 430 
mortoni, 430 
scotti, 430 
Arcteobia, 214, 225 
anticostiensis, 206, 225 
arctica, Aricia, 278 
Boreomysis, 579 
Castalia, 239 
Eteone, 234 
Eulalia, 238 
Nereis, 264 
Nicolea, 322 
Polynoé, 217 
Trophonia, 290, 291 
arcticus, Herpyllobius, 221, 223, 227 
Rangifer, 424 
arcuatus, Bolbelasmus, 97, 98, 99 (fig.), 
100, 114 (map) 
Bolboceras, 97, 98 
Kolbeus, 97, 98 
Ardynomys, 434 
arenarum, Ogmophis, 402 
Arenicola, 300 
glacialis, 205, 208, 300, 301 (fig.) 
Arenicolidae, 208 (table), 213, 300, 301 
(figs.) 
arge, Upeneoides, 518 
Upeneus, 500 (table), 502 (tables), 
503, 504 (table), 505 (tables), 
506, 507, 508, 518 
argentatus, Pleonomus, 613 
Aricia arctica, 278 
Ariciidae, 278 
arion, Amphiprion, 199 
arisan4s, Taiwanathous, 617 


636 


armatus, Idathyrsus, 
(fig.), 308 
armiger, Lumbricus, 278 
Scoloplos, 207, 278, 283 (fig.) 
armigera, Melania, 363 
armillaris, Nereis, 254 
Syllis, 254 
Arnett, Ross H., Jr.; A review of the 
beetle family Cephaloidae, 155 
Beetles of the oedemerid genus 
Vasaces Champion, 87 
Supplement and corrections of J. A. 
Hyslop’s Genotypes of the elaterid 
beetles of the world, 599-632 
Artiodactyla, 425, 432 
Asabellides, 315, 318 
orientalis, 318 
sibirica, 208, 318, 319 (fig.) 
Asclerini, 88 
Ascoliocerus, 603 
asinus, Equus, 423 
aspirans, Melania, 380 
Stenomelania, 380 
Asteidae, 560 
asymmetricus, Upeneus, 500 (table), 501, 
502 (tables) 503, 504 (table), 505 
(tables), 5/1, 512 (table) 
Athoina, 603 
Athous, 601, 603 
hirtus, 615 
revelieri, 603 
Athyreus ferrugineus, 102 
Atoloderma, 603 
atra, Doryssa, 367 
Atrichopogon, 66 
Athyreus (Bradicinetus) ferrugineus, 102 
serratus, 113 
(Bradicinetus) serratus, 113 
Aulacon, 603 
Aulacostoma, 375 
auratus, Mullus, 528, 529 
auriculata, Polycelis, 171, 172 
australasiae, Bolboceras, 101 
australis, Dennyus, 533, 534, 536, 537, 
540 (fig.), 549 
Eucopia, 578, 579 
Geckobia, 15 
austratica, Patricia, 614 
Autolytus, 241, 242, 245 (key), 249 
alexandri, 207, 242, 244, 245, 246 
cornutus, 244, 245, 250, 252 
fallax, 205, 207, 242, 244, 246, 247, 
250, 251 (fig.) 
incertus, 250 
longisetosus, 242, 247, 250 
ornatus, 244 
prismaticus, 207, 242, 244, 245, 246, 
247, 249, 251 (fig.) 
prolifer, 244, 245, 247, 249, 250 
trilineatus, 250 
verrilli, 246 
Aylacostoma, 367, 375, 376 
(Verena) crenocarina, 377, 378 
labrum, 375, 376 
Hemisinus) lineolatum, 376 
ruginosum, 377 


205, 208, 301 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 103 


(Aylacostoma) scalare, 376 
tuberculatum, 375, 377 
(Longiverena) tuberculatum, 377 


baccata, Brotia, 383 
badia, Antinoé, 216 
bailleti, Oxytrema, 368 
bairdi, Elasmognathus, 489 
Tapirus, 490 
Tapirus (Elasmognathus), 489 
bairdii, Elasmognathus, 488, 489 
Tapirella, 490 
Tapirus, 465, 466, 467, 468, 469, 
473, 478, 482 (fig.), 483 (fig.), 488, 
489, 490, 493 (map), 495 (map) 
Tapirus (Tapirella), 489 
bakeri, Hypnoidus, 610 
Balanocochlis, 381 
glans, 381 
balonnensis, Plotiopsis, 378 
bandi, Mullus, 516 
Banner, Albert H.; A supplement to 
W. M. Tattersall’s Review of the 
Mysidacea of the United States 
National Museum, 575 
barbata, Eteone, 206, 233 
Eteone (Mysta), 233 
Eunoé, 219 
Mysta, 233 
bartschi, Callocalia inexpectata, 543 
basalis, Chalcites, 594 
Hypnoidus, 603 
Basistoma, 376 
Bathanalia, 371, 374 
howesi, 374 
bathygenus, Parictis, 421 
Bathygenys, 434 
alfa, 430 
Bdellocephala, 174 
beckae, Stilobezzia, 59, 60, 61 
Stilobezzia (Stilobezzia), 
(fig.), 82 
bedeli, Blaiseus, 604 
beetle family Cephaloidae, A review 
of the, 155 
beetles, Chrysomelid, of the genus 
Strabala Chevrolat, 121 
Of the oedemerid genus Vasaces 
Champion, 87 
Secarabaeid, of the genus Brady- 
cinetulus, and closely related 
genera in the United States, 95 
Beezia stecki, 138 
behnii, Hemisinus, 375 
belaque, Upeneoides, 513 
Bembiciidae, 462 
Bembicium, 462 
bensasi, Mullus, 509 
Upeneus, 500 (table), 501, 502 
(tables), 503, 504 (table,) 505 
(tables), 506, 508, 509, 511 (table) 
Berger, Andrew J.; On the anatomy 
and relationships of glossy cuck- 
oos of the genera Chrysociccyx, 
Lampromorpha, and Chalcites, 
585-597 


69, 72 


INDEX 


Betarmon, 603 
quadrivittatus, 611 
Betarmou, 603 
bethaniensis, Photuris, 36 
Bezzia elegantulus, 62 
bicinctus, Amphiprion, 190, 192, 194, 
198, 199 
bicolor, Actinicola, 191 
Amphiprion, 191 
Cephaloon, 157 (map), 159, 160 
Stilobezzia, 59, 60, 61, 71 
Stilobezzia (Stilobezzia), 
(fig.) 
bifasciata, Wetmorella philippina, 440, 
441, 442 (fig.), 443 
bifasciatus, Amphiprion, 193 
Anthias, 194 
Prochilus, 193, 194 
bigeminatus, Elater, 602 
bilineatus, Cosmesus, 601 
birmanicus, Megapenthes, 619 
bisbistillatus, Fulvius, 627 (fig.), 631 
bispinus, Ambrysus, 3 (fig.), 4, 6 
bitaeniatus, Upeneus, 516 
bivittatus, Upeneus, 513 
Blaberus craniifer, 46 
Blaiseus, 604 
bedeli, 604 
Blake, Doris Holmes; The chrysomelid 
beetles of the genus Strabala 
Chevrolat, 121 
Blatta, 41 
livens, 40, 41 
livida, 40, 41 
maderae, 45 
surinamensis, 45 
Blattella, 53 
blomstrandi, Eusyllis, 207, 243 (fig.), 
260 
boholensis, Amphiprion, 198 
Boidae, 402 
Bolbelasmus, 95, 97 (key) 
arcuatus, 97, 98, 99 (fig.), 100, 114 


(map) 

gations, 97, 99 (fig.) 

hornii, 97, 98, 99 (fig.), 100, 115 
(map) 

minor, 97, 98, 99 (fig.), 100, 115 
(map) 

unicorne, 97 

Bolboceras, 95, 96, 101 

angulus, 95, 117 

arcuatus, 97, 98 

australasiae, 101 

coreanus, 97 

ferrugineus, 102 

(Amechamus) ferrugineus, 102 

fossatus, 103 

(Amechamus) fossatus, 103 

hornii, 100 

lecontei, 102 

minor, 100 

quadridens, 101 

schaefferi, 117, 119 

(Amechamus) serratus, 113 

serratus peninsularis, 116 


PE) RZ 


637 


Bolbocerastes, 95, 97, 99 (figs.), 105 
(key), 113 

imperialis, 99 (fig.), 106, 109, 112, 
113, 114 (map), 116 

imperialis kansanus, 106, 112, 115 
(map) | 

peninsularis, 98 (fig.), 105, 112, 


114 (map), 116 
105, 106, 109, 


regalis, 99 (fig.), 
112, 115 (map) 
serratus, 99 (fig.), 106, 107, 108, 
109, 112, 113, 115 (map) 
Bolboceratini, 96 (key) 
Bolbocerosoma, 95, 96 
Bolborhombus, 95, 97, 99 (figs.), 116, 
117 (key) 
angulus, 99 (fig.), 
TT a9 
parvul, 99 (fig.), 115 (map), 117, 
118, 119 


schaefferi, 99 (fig.), 114 (map), 
117, 119 
boliviana, Anischia, 603 
Boltenia echinata, 276 
iia Neti ea 168, 171 (fig.), 172, 
182 


115 (map), 


borealis, Micropternodus, 430 
Mystides, 206, 232, 237 
Nicomache lumbricalis, 305 
Petaloproctus tenuis, 306, 307 

Boreomysinae, 576 

Boreomysis, 579, 580 
arctica, 579 
californica, 579 
kineaidi, 577, 580 
microps, 576, 579 
tregonboffi, 579 
tridens, 579 

Bos depressicornis, 424 
frontalis, 424 
gaurus, 424 
indicus, 424 

Bothriodon, 433 
americanus, 430 

Bothriodontidae, 432 

Bourguignatia, 372 
imperialis, 372 

Bovidae, 424 

brachycephala, Nephthys, 271, 272 
Polydora, 280, 281 

Brachyfulvius, 621 
chapini, 622, 623 (fig.), 626 (fig.) 

Brachygonus, 604 
megerlei, 604 

brachyodon, Megalagus, 416, 430 

brachyura, Chaetura brachyura, 544 

Brada, 288, 290 (key) 
granosa, 292 
granulata, 290, 292 
inhabilis, 207, 290, 292, 297 (fig.) 
pilosa, 290, 291 
rugosa, 290, 291, 292 
setosa, 290, 291 
sublaevis, 292 
villosa, 205, 207, 290 

Bradycellus, 101 


(fig.) 


638 


Bradycinetulus, 95, 96, 97, 99 (figs.), 
101, 102 (key) 
ferrugineus, 96, 99 (fig.), 102, 103, 
104, 105, 115 (map) 
fossator, 103 
fossatus, 99 (fig.), 102, 103, 105, 
114 (map) 
hornii, 100 
rex, 99 (fig.), 102, 103, 114 (map) 
Scarabaeid beetles of the genus, 95 
serratus, 113 
Bradycinetus, 95, 101 
carinatus, 117, 119 
ferrugineus, 102 
fossatus, 103 
hornii, 95, 97, 100 
minor, 95, 97, 100 
serratus, 113 
serratus peninsularis, 116 
brahminus, Drasterius, 615 
Brania sp., 256 
brasiliensis, Mallerius, 612 
Monohelea, 137, 145 
Monohelea (Monohelea), 145 
Tapirus, 481, 485 
Tapirus pinchaque, 479 
Tapirus roulinii, 479 
braznikovi, Ampharete goési, 317 
brementi, Amyadenium, 177 
Dendrocoelopsis, 178 
breviata, Ammotrypane, 205, 208, 295, 
297 (fig.) 


brevicapitis, Dennyus, 535, 540 (fig.), 544 
brevicauda, Scalibregma, 293, 294 
Brevicerus, 604 

cylindricus, 604 
brevicollis, Creoleptus, 605 
brevicoma, Cistenides, 312 

Pectinaria, 312 

Pectinaria (Cistenaria), 312 
brevipennis, Hapithus, 24, 25 
brevis, Cubaedomus, 376 

Travisia, 298 
Brontotherioidea, 423, 432, 433 
Brotella, 383 
Brotia, 381, 382, 387 

baccata, 383 

pagodula, 382 
bruneri, Dennyus, 533, 534, 536, 537, 

588, 540 (fig.), 542 

Nitzschia, 538 

rn oes Dennyus, 535, 541 (fig.), 
48 


brunneitorques, Chaetura rutila, 548 
Bruyantius, 604 
capensis, 604 
buenoi, Ambrysus, 4 
Buffeventius, 604 
lividus, 604 
bulbosa, Oxytrema, 361 
bulla, Stilobezzia, 59, 61, 74, 83, 84 
Stilobezzia (Stilobezzia), 73 (fig.) 
74, 84 
burkei, Paleolagus, 416 
buxroni, Roggeveenia, 616 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 102 


Bythoceras, 373 
iridescens, 373 


caballus, Equus, 423 
Cacomantis, 586 
Cadiohypnus, 604 
caeca, Nephthys, 270 
Nereis, 270 
Caenopus, 424, 432 
mitis, 425, 430 
caerulea, Coua, 586 
caeruleus, Upeneoides, 516 
Caesaromysides liguriae, 557, 581 
Caesaromysis, 581 
sp., 577 
vanclevei, 577, 581 
Calamagras sp., 402 
californica, Boreomysis, 579 
Eteone, 234 
Calliostoma, 458 
Callocalia inexpectata bartschi, 543 
sp., 543 
Calostirus, 604 
cambodiensis, Corymbites, 618 
Camelidae, 426, 433 
Campsosdernus, 604 
Campsosternss, 604 
Campsosternus, 604 
camtschatica, Paralithodes, 224 
cancellata, Thiara, 378, 379 
Thiara (Setaeara), 379 
candezei, Mopleonus, 613 
Candezella, 604 
horvathi, 604 
Canidae, 416, 420, 423 
Caninae, 420 
Canis latrans, 423 
nubilus, 423 
canorus, Cuculus, 586 
Canyon Ferry Reservoir area, Pre- 
liminary analysis of the fossil 
vertebrates of, 395 
capensis, Bruyantius, 604 
Cardiotarsus, 605 
Longithorax, 581 
capitata, Capitella, 208, 297 (fig.), 298 
Glycera, 207, 272, 277 (fig.) 
capitatus, Lumbricus, 298 
Capitella, 298 
capitata, 208, 297 (fig.), 298 
capitata antarctica, 298 
giardi, 299 
Capitellidae, 208 (table), 
(figs.), 298 
Capitellides, 299 
caprius, Lampromorpha, 585, 586, 587, 
588, 589, 590, 591, 592, 593 (fig.), 595 
capsula habei, Littorina, 450 
Cardiohpynus, 604 
Cardiohypnus, 604 
mirabilis, 604 
Cardiophorus, 605 
florentini, 615 
schusteri, 611 
vitalisi, 613 


213, 297 


INDEX 


Cardiotarsus, 605 
capensis, 605 
Cariblatta sp., 43 
caribou, Rangifer, 424 
carinata, Mudalia, 361 
Nicomache, 305 
carinatus, Bradycinetus, 117, 119 
carnea, _Travisia 205, 208, 296, 297 


(fig.) 
Carnivora, 416, 432 
Cartwright, O. L.; Scarabaeid beetles 
of the genus Bradycinetulus and 
closely related genera in the 
United States, 95 
Carriker, M. A., Jr.; Studies in Neo- 
tropical Mallophaga, XI; Bird 
lice of the suborder Amblycera, 
genus Dennyus Neumann, 533 
Carvalho, José C. M.; Neotropical 
Miridae, LXIV; New bugs of the 
subfamily Cylapinae (Hemip- 
tera), 621 
Castalia, 239 
aphroditoides, 206, 239, 243 (fig.) 
arctica, 239 
fabricii, 239 
multipapillata, 239 
castaneus, Fulvius, 627 (fig.), 629 (fig.), 
630 


Castoridae, 408 
Catelanus, 605 

catenaria, Oxytrema, 360 
Cattleya sp., 632 

speciosissima, 632 
caudacutus, Hirundapis 
543 
a aaa penn 207, 280, 281, 283 
& 


caudacutus, 


cavernicola, Phagocata, 563, 565 (fig.) 
Cavicoxum, 605 
monstrosum, 605 
cecillei, Litorina, 456 
celebensis, Chesotraxus, 605 
celer, Mesohippus, 430 
Cenchrites, 462 
Cephalaon, 158 
Cephaloidae, 155, 156, 157 (map), 158, 
159 (key) 
A review of the beetle family, 155 
Cephaloon, 155, 156, 158, 160 (key) 
bicolor, 157 (map), 159, 160 
feptymeles: 156, 157 (map), 158, 
16 
pacificum, 157 (map), 159, 160 
pallens, 157 (map), 158, 160, 161 
pallens cinctipennie, 161 
pallens koltzei, 161 
pallens maculicolle, 161 
pallens picticolle, 161 
sakurae, 161 
tenuicorne, 157 (map), 158, 160 
ungulare, 157 (map), 158, 159, 160 
vandykei, 157 (map), 160 
variabile, 158 
variabilis, 157 (map), 158, 159, 161 
variabilis tristiculus, 161 


639 


cephus, Scarabaeus, 101 
Ceratolophus antennalis, 68 
diversus, 76 
copiosus, 138 
leucopeza, 138 
nebulosus, 151 
politus, 138 
viridis, 84 
Ceratopogon albitarsis, 138 
antennalis, 68 
copiosus, 137, 138 
diversus, 76 
leucopeza, 138 
maculipennis, 135, 140 
nebulosa, 135 
nebulosus, 151 
ochraceus, 63 
pictus, 75 
politus, 138 
sequax, 135 
viridis, 84 
Cercimelania, 366 
Cerithiidae, 357, 358 
Ceroleptus, 605 
brevicollis, 605 
Ceropectus, 605 
Cervidae, 423 
chadronensis, Hypertragulus, 430 
Chaetosyllis, 241, 252, 253, 255 
oerstedi, 253 
Chaetozone, 286, 287 
setosa, 207, 287, 297 (fig.) 
Chaetura brachyura brachyura, 544 
chapmani viridipennis, 545 
griseiventris, 539 
griseiventris phaseopygus, 539 
pelagica, 5 
richmondi, 537 
rutila brunneitorques, 548 
spinicauda aetherodroma, 539, 542 
spinicauda fumosa, 539, 542 
spinicauda spinicauda, 542 
Chalaraspidum alatum, 577 
Chalcites, 585, 589, 590, 591, 592, 593 
basalis, 594 
lucidus, 586, 587, 591, 592, 594, 595 
Chalcites-Chrysococcyx group, 594 
Chalcococcyx, 585, 586 
Chaleolepidinus, 605 
Chalcolepidius, 605 
challengeri, Lophogaster, 577 
chalybea, Progne chalybea, 549 
chapini, Brachyfulvius, 622, 623 (fig.), 
626 (fig.) 
chapmani, Panamenia, 555 
Scutops, 556 
Chatanayus, 605 
chattoni, Amyadenium, 177 
Dendrocoelopsis, 178 
Cheilinus, 439 
fasciatus, 439 
chelonyx, Mesoreodon, 425 
Chesotraxus, 605 
celebensis, 605 
Chiagosnius, 605 
chinensis, Melanoides, 380 


640 


Chloraemidae, 288 
Chone, 334, 337, 338 (key) 
dunéri, 209, 338, 339, 341 (fig.) 
gracilis, 338 
ce, RE CELT 209, 338, 341 
g. 
chopardi, Ectobius livens, 44 
Chorisoneura texensis, 42 
chrysargyrus, Amphiprion, 198 
Chrysis sp., 32 
Chrysococcyx, 585, 589, 590, 591, 592, 
594, 595 
cupreus, 585, 586, 587, 588 (fig.), 
589 (fig.), 590, 591, 592, 594, 595 
flavigularis, 593 
chrysogaster, Amphiprion, 189, 192, 193 
chrysopterus, Amphiprion, 198 
Amphiprion ephippium, 195 
Chusquea, 475 
ciliata, Gattyana, 205, 206, 226, 228 
Nephthys, 268, 270 
Nephtys, 207, 267, 269 (fig.), 270 
Nepthys, 270 
Mens 268, 270 
Cinchacus, 469 
cincinnata, Amphitrite, 327 
eee Thelepus, 209, 327, 33) 


g. 
cinctipennie, Cephaloon pallens, 161 
cinerea, Eteone, 234 

Nauphoeta, 40, 46, 51 
cingulatus, Cirratulus, 286 
Colodon, 430 
Circeis spirillum, 344 
circinnatus, Thelepus, 327 
cirrata, Ampharete, 316 
Amphitrite, 209, 319 (fig.), 321 
Enipo, 225, 305 
Lepidonote, 220 
Polynoé, 220 
Cirratuligae’ 207 (table), 212, 285, 28t 
(key), 297 (figs.) 
Cirratulus, 286 
cingulatus, 286 
cirratus, 207, 286, 297 (fig.) 
robustus, 286 
spectabilis, 286 
cirratus, Cirratulus, 207, 286, 297 (fig.) 
Lumbricus, 286 
cirrhosa, Aphrodita, 226 
Gattyana, 227 
cirrosa, Gattyana, 206, 226, 229 (fig.) 
Nychia, 226 
Cistenides, 311 
brevicoma, 312 
granulata, 312 
hyperborea, 314 
Citellus, 434 
citrina, Phyllodoce, 236 
Clamator, 594 
clarki, Eunoé, 205, 206, 216, 217 
clarckii, Amphiprion, 198 
Anthias, 198 
Cleniocerus, 606 
Clinopternodus, 431, 433 
gracilis, 430 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 103 


Clymene (Praxillella) praetermissa, 303 
coarctata, Phagocata, 167 
coarctatus, Hyas, 224, 324 
Coccyzus, 590, 591, 592 
erythropthalmus, 588, 589 
cockroaches, Distribution, general bi- 
onomics, and recognition char- 
acters of, recently established in 
the United States, 39 
coeca, Nephthys, 268 
Colaulon, 606 
rectangularis, 606 
collaris, Typosyllis, 260, 261 
Colodon, 482, 433 
accidentalis, 430 
cingulatus, 430 
colombianus, Tapirus terrestris, 486, 490, 
493 (map), 495 (map) 
eg haste Strabala, 125 (fig.), 127, 
133 
comalensis, Oxytrema, 360 
compacta, Pionosyllis, 205, 207, 243 
(fig.), 262 
concinna, Nullarborica, 613 
concolor, Felis, 424, 475 
Conobajulus, 606 
ugiensis, 606 
consanguineus, Photinus, 36 
consolidata, Dorssa, 367 
conspersa, Adelocera, 607 
convexuni, Yezodina, 619 
coomani, Heabinh, 610 
copiosa, Schizohelea, 138 
copiosus, Ceratolophus, 138 
Ceratopogon, 1387, 138 
coquilletti, Stilobezzia, 58, 60, 61, 75 
ati ea (Stilobezzia), 72 (fig.), 


7 
Coracinus, 194 
intermedius, 194 
vittatus, 194 
Corcovadocola, 622 
coreanus, Bolboceras, 97 
Coresus, 600, 601, 606 
cornuta, Ehlersia, 253 
Syllis, 205, 207, 248 (fig.), 252, 254 
Syllis (Ehlersia), 253 
cornutus, Autolytus, 244, 245, 250, 252 
coronarius, Tectarius, 461 
coronata, Lavigeria, 372 
Polycelis, 171, 172 
corvinus, Pachychilus, 365, 366 
Coryleus, 606 
desruisseauxi, 606 
Corymbites cambodiensis, 618 
gratus, 613 
sulphuripennis, 616 
tenuicornis, 614 
tessellatus, 600 
Cosmesus, 601 
bilineatus, 601 
subtilis, 601 
cosmius, Ambrysus, 6 
costaricensis, Strabala acuminata, 
(fig.), 127, 129 
costatus, Vasaces, 87, 89, 90 


125 


INDEX 


costula, Antimelania, 382 
Coua, 590, 595 

caerulea, 586 
Couroupita guianensis, 491 
craniifer, Blaberus, 46 
crassa, EKurycaelon, 362 
crassicornis, Sabella, 209, 333 (fig.), 334 
crassum, Archaeotherium, 430 
Cratonychus, 606 
crawfordensis, Hypertragulus, 430 
crawshayi, Melanoides, 371 
Sepererine, Aylacostoma (Verena), 377, 

8 


Melania, 377 
Cricedon, 412 
Cricetidae, 410 
cricetodontoides, Eumys, 410, 411 (fig.) 
Cricket parasite of the genus Rhopal- 
osoma, Notes on the biology and 
immature stages of a, 19 
cristata, Eurycaelon, 362 
Crotophaga, 590, 595 
sulcirostris, 590 
crucifer, Elater, 602 
cruentus, Hyaenodon, 430 
Crypnoidus, 606 
Cryptohypnus, 606 
micros, 615 
Cryptypnus, 606 
Csikia, 607 
dimatoides, 607 
Ctenicera, 600, 607 
pupieri, 615 
(Malloea) sjaelandica, 600 
Ctenicerus, 607 
Cteniocerus, 607 
Ctenocera, 607 
Ctenonychus, 607 
Ctenonyphus, 607 
Cubaedomus, 366, 376, 387 
brevis, 376 
Cuculus, 585, 594 
canorus, 586 
culbertsoni, Merycoidodon, 425 
cumingi, Echininus, 460 
Trochus, 458 
cumingii, Pachychilus, 366 
cupreus, Chrysococcyx, 585, 586, 587, 
588 (fig.), 589 (fig.), 590, 591, 592, 
594, 595 
Curtisius, 607 
Cyamops, 551, 555, ial 557, 558 (key) 
imitata, 558, 559 
nebulosa, 558 
cyaneus, Psephus, 611 
Cyclopidius, 426 
lullianus, 426 
simus, 426 
cyclops, Scarabaeus, 101 
Cylapinae, 621, 622 
cylindricus, Brevicerus, 604 
Lincydrus, 612 
Cylindrodon, 431, 433 
fontis, 430 
Cypseloides fumigatus, 547 
cypsiurus, Dennyus, 534, 535 


641 


Dactylosimus, 607 
dorsalis, 607 
dakotensis, Ictops, 430 
Parictis, 420, 430 
dalli, Pachychilus (Pilsbrychilus), 367 
Dalopius, 607, 608 
damoni, Paramelania, 371 
Danosoma, 607 
Daphoenidae, 420 
Daphoenocyon, 420, 421, 432, 433 
dodgei, 419, 421 (fig.), 430 
Daphoenus, 420 
davisi, Acanthomysis, 582 
de bojer, Amphiprion, 198 
Deinictis, 433 
fortis, 430 
Delox, 607 
Dendrocelopsis spinosipenis, 176 
Dendrocoelidae, 174, 568 
Dendrocoelopsis, 174, 176, 177, 178 
ean 165, 168, 177, 178, 181 


brementi, 178 

chattoni, 178 

garmieri, 178 

piriformis, 174, 177 (fig.), 178, 180 

spinosipenis, 177, 178 

vaginata, 178 

vaginatus, 177 

vandeli, 178 

Dendrocoelum, 178 
nausicaae, 179 
Dennyus, 533, 535 (key), 549, 534 

australis, gen, 534, 536, 537, 540 
(fig.), 549 

brevicapitis, 535, 540 (fig.), 544 

bruneri, 533, 534, 536, 537, 538, 
540 (figs.), 542 

brunneitorques, 535, 541 (fig.), 548 

cypsiurus, 534, 535 

distinctus, 534, 548 

dubius, 533, 536, 537, 538, 542 

femuralis, 533 

francicus, 534, 535 

gossei, 534, 536, 543, 547 

hirundinidis, 534, 535 

intonsus, 535, 540 (fig.), 645 

latifrons, 533 

limbus, 536, 538, 540 (fig.), 542 

major, 534, 535, 643 

meridionalis, 534, 536, 539, 540 
(fig.), 542 


piageti, 533 

richmondi, 533, 535, 537 

rotundocapitis, 535, 541 (fig.), 548, 
549 


similis, 534, 535, 541 (fig.), 549 

spiniger, 533, 535 

spininotus, 536, 541 (fig.), 547 
depressa, Eteone, 235 

Eunoé, 217, 218 
depressicornis, Bos, 424 
descarpentriesi, Practapyrus, 615 
deshayesiana, Oxytrema, 361 


642 


Desmatolagus, 432, 433, 434 
dicei, 430 
desruisseauxi, Coryleus, 606 
Dexiospira spirillum, 344 
Diacanthus, 608 
guttatus, 610 
Diadelops, 560 
dicei, Desmatolagus, 430 
Dichonychus, 608 
Dicrepidius, 608 
Dicronychus, 608 
serraticornis, 609 
Didelphiade, 402 
Dido, 600 
macrocera, 600 
dilatata, Anculosa, 363 
Littorina, 451 
dilaticollis, Eupsephus, 609 
dilatus, Ambrysus, 6 
dimatoides, Csikia, 607 
Diocarphus, 608 
Diopsosoma, 555, 556 
prima, 556 
Diphyaulon, 608 
Diploccnus, 608 
Diploconus, 608 
ornatus, 616 
spiloderus, 615 
Diploptera, 51, 52 
discors, Nephthys, 270 
Nephtys, 207, 267, 269 (fig.), 270 
Discrepeidius, 608 
Disonycha, 122, 124, 126 
rufa, 122 
dispar, Exogone, 207, 243 (fig.), 258, 
259 


Heteromeryx, 430 
Paedophylax, 259 

distinctus, Dennyus, 534, 543 

diversa, Hortomyia, 76 
Johannseniella, 76 
Stelobezzia, 59, 60, 61, 76 
Stilobezzia (Stilobezzia), 73 (fig.), 

76, 79 

diversus, Ceratolophus, 76 
Ceratopogon, 76 

Dodecaceria, 285 

dodge cer one 419, 421 (fig.), 


Dolopus, 608 
dolosus, Elastrus, 602 
Domenephus, 608 
dominicensis, Strabala, 121 
Domnina, 431, 432, 433, 434 
thompsoni, 430 
doreas, Gazella, 424 
dorsalis, Dactylosimus, 607 
dorsofasciata, Palpomyia, 62 
Dorygonus alluaudi, 617 
Doryssa, 367, 377 
atra, 367 
consolidata, 367 
Drachylis, 158 
dowi, Elasmognathus, 489 
Tapirella, 490 
Tapirus, 490, 492 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 103 


dowii, Elasmognathus, 489 
Tapirus, 480, 484, 490, 491 
Tapirus bairdii, 491 

Drachylis simulans, 158, 160 

drakei, Pachychilus, 360 

Drasterius, 608 
brahminus, 615 

Dresterius, 608 

Drosophilidae, 553 

dubius, Dennyus, 533, 536, 537, 538, 542 
Mullus, 498, 515 
Nitzschia, 537 
Upeneoides, 514 

dufourii, Melanopsis, 359 

dunéri, Chone, 209, 339, 341 (fig.) 


eburnea, Polycelis, 172 
echinata, Boltenia, 276 
Echinella, 449, 458, 461 
Kchininus, 449, 451, 452, 454, 455 (fig.), 
458, 459, 461 
cumingi, 460 
nodulosus, 449, 451, 452, 453 (fig.), 
454, 455 (fig.), 457 (fig.), 468 
Ectobius, 40 
finoti, 41 
lapponicus, 41, 45 
livens, 39, 40, 47 (fig.) 
livens chopardi, 44 
livens minor, 44 
lividus, 40, 41 
panzeri, 44 
perspicillaris, 41 
siculus, 41 
sp., 41 
ecuadorensis, Tapirus, 479, 481, 485 
Edgaria, 372 
paucicostata, 372 
Ehlersia cornuta, 253 
Elasmognathus, 488 
bairdi, 489 
bairdii, 488, 489 
dowi, 489 
dowii, 489 
Elastrus dolosus, 602 
Elater aeneus, 608 
bigeminatus, 602 
crucifer, 602 
fascicularis, 615 
ferrugineus, 612 
fulvipes, 612 
granulipennis, 618 
hirtus, 615 
lepidotus, 616 
linearis, 609 
lineatus, 610 
marginatus, 607 
megerlei, 604 
murinus, 602 
nobilis, 601 
obesus, 608 
obscuripes, 605 
obscurus, 606 
ovalis, 602 
pallidulus, 602 
pectinicornis, 606, 607 


INDEX 


purpureus, 614 
riparius, 606, 610 
sjaelandicus, 600 
tessellatus, 600 
thoracicus, 605 
vittatus, 603 
Elaterid beetles of the world, Supple- 
ment and corrections to J. A. Hy- 


643 


flava, 206, 233, 235, 237 (fig.) 
lentigera, 235 

longa, 206, 233, 234, 237 (fig.) 
Sarsi, 235 

sp. (larvae), 236 
spetsbergensis, 206, 233, 235 
striata, 233 

tuberculata, 234 


slop’s genotypes of the, 599-619 Eucanthus, 95, 97 


Elateridae, 599 
elegans, Eumys, 410 
elegantula, Eukraiohelea, 62 
Parabezzia, 62 
Probezzia, 62 
Stilobezzia, 58, 60, 61 
Syabeaais (Eukraiohelea), 62, 72 


g. 

elegantulus, Bezzia, 62 
Ellipstoma, 363 

gibbosa, 363 

zonalis, 363 
elongata, Polycelis, 172 
elongatus, Vasaces, 89, 91 (fig.), 93 
Emmericia, 386 
Emmericiinae, 386 
Endicronychus, 609 
enigmaticus, Tapirus, 471, 493 (map) 
Enipo, 214, 225 

cirrata, 225, 305 

gracilis, 206, 225, 305 
Entelodontidae, 432, 435 
Eomyidae, 409, 432 
Kotylopus, 433, 434 

reedi, 430 
Ephamillus, 158, 159 

variabilis tristiculus, 161 
egg Amphiprion, 190, 192, 199. 

201 


Lutjanus, 188, 199 
Prochilos, 199 
episcopalis, Antimelania, 382 
Melania, 382 
Equidae, 421, 423 
Equus asinus, 423 
caballus, 423 
onager, 424 
prezevalskii, 424 
Ereutho smitti, 328 
erinaceus, Sphaerosyllis, 207, 243 (fig.), 
255 
erythraeus, Lophogaster, 577 
Lophogaster typica, 577 
Erythropini, 576 
erythropthalmus, Coccyzus, 588, 589 
esakiana, Stilobezzia (Eukraiohelea), 61, 
62 


Espeletia, 475 
esperi, Fagotia, 359 
esulcatus, Leptomeryx, 426, 427 
Kteone, 231, 232, 233 (key), 237 (fig.) 
arctica, 234 
barbata, 206, 233 
(Mysta) barbata, 233 
californica, 234 
cinerea, 234 
depressa, 235 


Euchaetomera, 580 

tenuis, 580 
Euchaetomeropois, 580 

pacifica, 577, 580 
Euchone, 334, 339 

analis, 209, 339, 341 (fig.) 
Eucopia, 578 

australis, 578, 579 

grimaldii, 579 

hanseni, 479 

major, 579 

sculpticauda, 578 

unguiculata, 579 
Eucranta anticostiensis, 226 
Eudactylus, 618 

wapleri, 618 
eudendrocoeloides, Polycelis, 172 
Eudicrenychus, 609 
Eukraiohelea, 61, 62, 83 

africana, 61 

amnigena, 61 

elegantula, 62 

foyi, 61 

inusitata, 61 

versicolor, 61, 62 
Eulalia, 238 

arctica, 238 

minuta, 238 
Eumecomera, 88 
Eumida, 238 

minuta, 205, 206, 237 (fig.), 238 
Eumys, 410 (key) 

cricetodontoides, 410, 411 (fig.) 

elegans, 410 

exiguus, 411, 415 

latadens, 411, 412, 413 (fig.) 

spokanensis, 411, 413, 414 (fig.) 
Eunoa nodosa, 217 
Eunoé, 214, 216 (key) 

barbata, 219 

clarki, 205, 206, 216, 217 

depressa, 217, 218 

nasoe, 206, 216, 217, 219, 229 

(fig.) 


oerstedi, 206, 216, 218, 219, 229 
(fig.) 

Eupolynoé anticostiensis, 225 
Eupsephus, 609 

dilaticollis, 609 
eurantron, Polycelis, 172 
Eureum, 544 
Eurycaelon, 362 

anthonyi, 362 

crassa, 362 

cristata, 362 
Eurycotis floridana, 46 


644 


Eusyllis, 242, 259, 260 (key), 262 
blomstrandi, 207, 243 (fig.), 260 
magnifica, 205, 207, 260, 261 
monilicornis, 260 
phosphorea, 260, 261 
tubifex, 260 

Eutypomys, 432, 433 
thompsoni, 430 

evansi, Leptomeryx, 427 (table) 

Evarnella triannulata, 223 

excelsus, Tapirus, 468 

excisum, Gyrotoma, 361 

exiguus, Eumys, 411, 415 

eximium, Poébrotherium, 426, 430 

Exogone, 242, 257, 258 (ke y) 
dispar, 207, 243 (fig.), 258, 259 
gemmifera, 258 
lourei, 259 

naidina, 207, 243 (fig.), 258 

Exophthalmus, 609 

expallidus, Nyctor, 613 

extenuata, Harmothoé, 206, 220, 222 
Lagisca, 222 
Polynoé, 222 


Fabricia sabella, 299 
fabricii, Castalia, 239 
Sabella, 334 
Fagotia esperi, 359 
fairchildi, Photuris, 37 
fairmairei, Medonia, 616 
fallax, Autolytus, 205, 207, 242, 244, 
246, 247, 250, 251 (fig.) 
farctus, Scarabaeus, "101 
fasciata, Syllis, 205, 207, 243 (fig.) 
Syllis (Typosyllis), 254 
fasciatus, Cheilinus, 439 
fascicularis, Elater, 615 
fasciolata, Melanoides, 380 
fasciolatus, Upeneoides, 514 
fascipennis, Scutops, 555, 556 
Felidae, 4 
felina, Polycelis, 172 
Felis concolor, 424, 475 
femuralis, Dennyus, 533 
ferruginea, Altica, 121, 132 
Lactica, 123 
Strabala, 122, 125 (fig.), 127, 132 
ferrugineus, Amechanus, 102 
Athyreus, 102 
Athyreus (Bradicinetus), 102 
Bolboceras, 102 
Bolboceras (Amechamus), 102 
Bradycinetulus, 96, 99 (fig.), 102, 
103, 104, 105, 115 (map) 
Bradycinetus, 102 
Elater, 612 
Scarabaeus, 101, 102 
fiebrigi, Stilobezzia, 76 
Fijidoma, 383, 385, 386 
laddi, 383, 385 
maculata, 383, 385 
filicornis, Nereis, 284 
Spio, 207, 283 (fig.), 284 
filifer, Petaloproctus, 306, 307 
filifera, Maldane, 306 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 108 


finoti, Ectobius, 41 
firefly, lampyrid, Photuris bethaniensis, 
new, 35 
fish genus Wetmorella, Review of, with 
descriptions of new forms from 
the tropical Indo-Pacific, 439 
fishes, Indo-Pacific anemone, Review 
of the, genus Amphiprion, with 
deseriptions of two new species, 
Flabelligera, 288, 289 
affinis, 207, 289, 297 (fig.) 
infundibulariformis, 289 
infundibularis, 289 
infundibularum, 289 
Flabelligeridae, 207 (table), 212, 213, 
288 (key), 297 (figs.) 
flava, Eteone, 206, 233, 235, 237 (fig.) 
Nereis, 235 
flavangulus, Phedomenus,608 
flavigularis, Chrysococcyx, 593 
Fleutiauxellus, 609 
flexuosa, Pista, 226 
flies, Neararctic, of the family Peris- 
celidae (Diptera) and certain 
Anthomyzidae referred to the 
family, 551 
flinti, Phorticoides, 553, 556 
floccosa, Lagisca, 222 
Polynoé, 222 
florentini, Cardiophorus, 615 
ores Strabala rufa, 125 (fig.), 127, 
28 
floridana, Eurycotis, 46 
fluvialis, Io, 363, 364 
Fonticola, 166, 167 
fontis, Cylindrodon, 430 
foraminifera, Polynoé, 217 
forbesi, Travisia, 296 
forbesii, Travisia, 296 
Forficula livida, 41 
formosanus, Neolacon, 613 
fortis, Deinictis, 430 
fossator, Bradycinetulus, 103 
fossatus, Amechanus, 103 
Bradycinetulus, 99 (fig.), 102, 103, 
105, 114 (map) 
Bradycinetus, 103 
Bolboceras, 103 
Bolboceras (Amechamus), 103 
fossor, Sternaspis, 309 
foyi, EKukraiohelea, 61. 
fragilis, Lumbrinereis, 275 
Lumbrineris, 207, 275, 277 (fig.) 
Lumbriconereis, 275 
Lumbricus, 275 
francicus, Dennyus, 534, 535 
franciscorum, Neomysis, 582 
francki, Gnathodicrus, 610 
freethii, Potadoma, 370 
frenatus, Amphiprion, 180; 192, 199, 200 
Hemidactylus, 9, 11, 
frontalis, Bos, 424 
fuentesi, Pachychilus, 366 
fugax, Peratherium, 402 


INDEX 


fulvicollis, Pengamethes, 614 

Fulviini, 621, 622 

fulvipennis, Tenalomus, 618 

fulvipes, Elater, 612 

Fulvius albonotatus, 624 (fig.), 627 (fig.), 

629 (fig.) 

bisbistillatus, 627 (fig.), 631 
castaneous, 627 (fig.), 629 (fig.), 630 
imbecilus, 631 
lunulatus, 630 
ornatifrons, 626 (fig.), 631 
quadristillatus, 627 (fig.), 631 

fumigatus, Cypseloides, 547 

fumosa, Chaetura spinicauda, 539, 542 

funebris, Ambrysus, 1 

fusciventer, Amphiprion, 193 

fuscula, Stilobezzia, 58, 59, 61, 64 
se (Neostilobezzia), 64, 72 

g. 
Fusimorphus, 609 
Fusus, 364 


gallicus, Bolbelasmus, 97, 99 (fig.) 
Scarabaeus, 97 
Gamepenthes, 609 
Ganoxanthus, 609 
garmieri, Amyadenium, 177 
Dendrocoelopsis, 178 
Gattyana, 214, 226 (key) 
ciliata, 205, 206, 226, 228 
cirrhosa, 227 
cirrosa, 206, 226, 229 (fig.) 
gaurus, Bos, 424, 
Gazella dorcas, 424 
granti, 424 
leptoceros, 424 
soemmerringii, 424 
thomsonii, 424 
Geckobia, 9, 10, 15 
australis, 15 
hindustanica, 15 
keegani, 10 (fig.), 11 (fig.) 
philippinensis, 12, 13 (fz. ), 14 (fig.) 
simplex, 12 
sp., 9, 10 
texana, 15 
Geckobiella, 15 
texana, 9, 15, 16 (fig.), 17 (fig.) 
geismarianus, ’Nothocyon, 419 
gemmifera, Exogone, 258 
geniculata, Lithasia, 363 
Geococcyx, 590, 595 
geographica, Adelocera, 617 
seorgtenn, Sphaloplana, 565 (fig.), 56¢ 
567 (fig.), 569 (fig.) 
Georgicus, 609 
sanguinipennis, 609 
Geraniella, 610 
Geranus, 610 
giardi, Capitella, 299 
gibbosa, Ellipstoma, 363 
Pleurocera (Ellipstoma), 363 
gigantea, Melaenis lovéni, 214 
gigas, Gnathophausia, 578 


645 


gilva, Hartomyia, 65, 66 
glabrum, Aylacostoma, 375, 376 
glacialis, Arenicola, 205, 208, 300, 301 


(fig.) 
Trichobranchus, 209, 329, 331 (fig.) 
glans, Balanocochlis, 381 
Melania, 381 
glaphyrus, Glyptomelania, 366 
Pachychilus, 365 
glauca, Iphis, 611 
Lacais, 611 
Stilobezzia, 59, 60, 61, 77, 85 
Stilobezzia (Stilobezzia), 73 (fig.), 
77 


Glossophaga soricina soricina, 465 
Glycera, 272 
capitata, 207, 272, 277 (fig.) 
lapidum 273 
nana, 273 
setosa, 272 
Neuer (table), 213, 272, 277 


gs. 

Glycinde, 274 

wiréni, 205, 207, 274, 277 (fig.) 
Glyptomelania, 366 

glaphyrus, 366 
Glyptosaurus montanus, 401 
Gnathedicrus, 610 

francki, 610 
Gnathophausia, 578 

gigas, 578 

gracilis, 578 

ingens, 578 

zoca, 578 
goési, Ampharete, 308, 317 
Goniada nordmanni, 274 
Goniadidae, 207 (table), 213, 274, 277 


_ _ (figs.) 

Goniobasis, 359, 360, 361 
livescens, 361 
osculata, 261 

Gonodyrus, 610 
tarsalis, 610 

gossei, Dennyus, 534, 536, 543, 547 

gottschei, Melania, 370 

gracilis, Chone, 338 
Clinopternodus, 430 
Enipo, 206, 226, 305 
Gnathophausia, 578 
Polynoé, 225 
Proceraea, 250 

gradatus, Paleocastor, 408 

graffi, Proclea, 325 

graffi, Leaena, 325 
Proclea, 209, 325 

grandifolia, Heteronereis, 264, 265 

grandis, Lavigeria, 372 
Leuconhaea, 51 
Loxorhynchus, 299 

granifera, Melania, 379 
Tarebia, 379, 386 

graniperda, Melania, 370 

granosa, Brada, 292 

eranti, Gazella, 424 


646 


granulata, Brada, 290, 292 
Cistenides, 312 
Pectinaria, 208, 313 (fig.) 
Pectinaria (Cistenides), 311, 312 
Sabella, 312 
Serpula, 343 
granulatus, Spirorbis, 209, 341 (fig.) 
Spirorbis (Lacospira), 843 
granulipennis, Elater, 618 
graphium, Pachychilus, 366 
gratiosus, Agriotes (Ectinus), 618 
gratus, Corymbites, 613 
grebnitzkii, Archaeomysis, 580 
gregarius, Hesperocyon, 418 (fig.), 430 
grimaldii, Eucopia, 579 
griseiventris, Chaetura, 539 
groenlandica, Amphitrite, 209, 321, 322 
Anaitides, 236 
Ophelina, 295, 296 
Phyllodoce, 206, 236, 237 (fig.) 
Phyllodoce (Anaitides), 236 
Pista, 324 
Gromphadorhina, 51, 52 
laevigata, 52 
grubei, Ampharete, 316 
gryllodes, Orocharis, 24 
guianae, Monohelea, 137, 147 (fig.) 
Monohelea (Monohelea), 150 
guianae, Tapirus terrestris, 480, 481, 485 
guianensis, Couroupita, 491 
Gurney, Ashley B.; Distribution, general 
bionomics, and recognition char- 
aracters of two cockroaches re- 
cently established in the United 
States, 39 
Notes on the biology and immature 
stages of a cricket parasite of the 
genus Rhepalosoma, 19 
guttatipennis, Ambrysus, 6 
guttatus, Diacanthus, 610 
Selatosomus, 610 
Upeneoides, 509 
Gyrotoma, 361 
excisum, 361 
ovoideum, 361 
Gyrus, 610 


haematina, Strabala, 134 
hagruma, Littorina, 450 
Haltica, 122 
ambulans, 122, 123, 131 
rufa, 121, 122, 123, 127 
scutellaris, 123 
hamatus, Thelepus, 327 
Hamus, 451 
hanseni, Eucopia, 479 
Hapatesus, 610 
Hapithus agitator, 20, 21, 23, 24, 25, 31 
agitator agitator, 23 
agitator quadratus, 23, 24 
brevipennis, 24, 25 
sp., 21, 22, 23, 24; :25, 32 
Haploscloloplos alaskensis, 
Harmothoé, 214, 220 (key) 
anticostiensis, 225 
extenuata, 206, 220, 222 


278, 279 


PROCEEDINGS OF THE 


NATIONAL MUSEUM VOL, 1038 


imbricata, 206, 220, 229 (fig.) 
levis, 221 
multisetosa, 223 
nodosa, 217, 219 
propinqua, 222 
rarispina, 222 
rarispina propinqua, 222 
sarsi, 215 
(Antinoélla) sarsi, 215 
triannulata, 222, 223 
Harpagocryptus, 33 
Hartomyia antennalis, 68 
gilva, 65, 66 
lutea, 65, 66 
pallidiventris, 78 
picta, 75 
viridis, 84 
Hartomyis nebulosa, 151 
hastula, Melania, 380 
Haupathesus, 610 
hawaiensis, Lophogaster, 577 
haydeni, Paleolagus, 415 
hebes, Photuris, 36 
Heleidae, 57, 136, 141 
Heliconia mariae, 630 
Heliocis, 88 
Hemiblabera tenebricosa, 46 
Hemidactylus frenatus, 9, 11, 14 
leschenaulti, 12, 15 
Hemimitra, 369 
hemionus, Odocoileus, 428 
Hemiptera, 488 
Hemirhaphes, 610 
Hemirhipus, 610 
submetallicus, 609 
trilineatus, 605 
Hemirrhaphes, 610 
Hemisinus, 367, 375, 376 
behnii, 375 
lineolata, 375 
henriettae, Melania, 383 
Heptacodon, 433 
sp., 430 
Herpyllobius arcticus, 221, 223, 227 
Hershkovitz, Philip, Mammals of north- 
ern Colombia, preliminary report 
No. 7; (genus Tapirus), with a 
che he review of American 
species, 
Hesionidae, 306. table), 212, 239, 243 


gs 
Hesperocyon, 420, 432, 433 
gregarius, 418 (fig ly 430 
paterculus, 416, 417 (fig.), 430 
sp., 418 
heterochaeta, Syllis, 254 
Syllis (Ehlersia), 253 
Heteromeryx, 433, 434 
dispar, 430 
Heteronereis grandifolia, 264, 265 
Hevea sp., 630 
hieroglyphica, Monohelea, 135, 136, 137, 
139, 140, 141, 142, 143, 147 ‘(fig.) 
Monohelea (Monohelea), 140, 144, 
14 
hindustanica, Geckobia, 15 


INDEX 


Hippopotamus terrestris, 479 
hirtus, Athous, 615 
Elater, 615 
Hirundapis caudacutus caudacutus, 543 
hirundinidis, Dennyus, 534, 535 
Hoabinh, 610 
coomani, 610 
hoffmasteri, Speophila, 567 (fig.), 569 
(fig.), 570, 571 (fig.) 
Holmesiella anomala, 580 
Hoplophoneus, 433 
mentalis, 430 
o’harrai, 430 
robustus, 430 
horei, Tiphobia, 373, 374 
Horizoteichos, 610 
papuensis, 610 
hornii, Bolbelasmus, 97, 98, 99 (fig.), 
100, 115 (map) 
Bolboceras, 100 
Bradycinetulus, 100 
Bradycinetus, 95, 97, 100 
Kolbeus, 100 
horribilis, Ursus, 419, 424 
Hortomyia diversa, 76 
horvathi, Candezella, 604 
howesi, Bathanalia, 374 
Hua, 368 
hudsonica, Nepthys, 270 
hugeli, Paracrostoma, 383 
hungerfordi, Ambrysus, 6 
Hyaenodon, 434 
cruentus, 430 
montanus, 430 
Hyas coarctatus, 224, 324 
coarctatus alutaceus, 261 
Hydrobiidae, 386 
Hydrochaerus tapir, 479 
hyeroglyphica, Monohelea, 140 
Hyman, Libbie H.: North American 
triclad Turbellaria, XIII: three 
new cave planarians, 563 
Hypdonus, 610 
Hypeneus, 506 
vittatus, 516 
hyperborea, Cistenides, 314 
Pectinaria, 208, 313 (fig.) 
Pectinaria (Cistenides), 311, 314 
Hypertragulidae, 426, 432, 433 
Hypertragulus, 433, 434 
chadronensis, 430 
crawfordensis, 430 
Hypisodus, 433, 434 
paululus, 430 
Hypnoidus, 610 
bakeri, 610 
basalis, 603 
maritimus, 609 
Hypolithus, 610 
Hypolittus, 610 
hypostylus, Mesohippus, 
(table), 430 
Hyracodon, 432 
acridens, 430 
Hyracodontidae, 428 


421, 422 


647 


Ichnodes, 158 
Ictops, 431, 433 
acutidens, 403, 430 
dakotensis, 430 
thompsoni, 430 
Idanthyrsus, 308 
armatus, 205, 208, 301 (fig.), 308 
ornamentatus, 308 
Idiotarmon, 611 
ignobilis, Melanoides, 371 
Ijimia, 172 
imbecilis, Fulvius, 631 
imbricata, Aphrodita, 220 
Harmothoé, 206, 220, 229 (fig.) 
tmitata, Cyamops, 558, 559 
impatiens, Lagisea, 222, 223 
imperialis, Bolbocerastes, 99 (fig.), 106, 
109, 112, 113, 114 (map), 116 
imperialis, Bourguignatia, 372 
incertus, Autolytus, 250 
indefinata, Melanoides, 380 
indicus, Bos, 424 
Tapirus, 465, 469 
Tapirus (Acrocodia), 467, 489 
Indo-Pacific anemone fishes, genus Am- 
phiprion, with descriptions of 
two new species, Review of, 187 
inermis, Melania, 381 
inflatum, Scalibregma, 208, 293, 297 


(fig.) 

inflex, Paleogale, 430 
infundibulariformis, Flabelligera, 289 
infundibularis, FlabeHigera, 289 
infundibularum, Flabelligera, 289 
infundibuliformis, Chone, 209, 338, 341 


(fig.) 
infundibulum, Amphitrite, 340 
Myxicola, 209, 340, 341 (fig.) 
ingens, Gnathophausia, 578 
inhabile, Siphonostoma, 292 
inhabilis, Brada, 207, 290, 292, 297 


_(fig.) 
Insectivora, 403, 431 
insulare, Lophogaster subglaber, 577 
intermedia, Lactica, 123 
Pseudopotamilla, 337 
Strabala, 121, 122, 125 (fig.), 127, 
131, 133 
intermedius, Amphiprion, 194 
Coracinus, 194 
Lophogaster, 577 
Paleolagus, 415 
intonsus, Dennyus, 535, 540 (fig.) 646 
inusitata, Eukraiohelea, 61 
Inusitatomysis serrata, 581 
sp., 581 
Io, 363 
fluvialis, 363, 364 
spinosa, 364 
tenebrosa, 364 
Iphis glauca, 611 
Ipostirus, 611 
parumcostatus, 611 
iridescens, Bythoceras, 373 
irrorata, Littorina, 460 


648 


Ischnomyia, 557 
Ischyromyidae, 408 
Ischyromys, 431, 432, 433, 434 
pliacus, 408, 430 
islandica, Polynoé, 217, 218 
Sternaspis, 309 


PROCEEDINGS OF 


jacquetiana, Melania, 368 
Oxytrema, 368 
jamaicensis, Strabala ambulans, 
(fig.), 127, 131 
japonica, Terebellides stroemii, 330 
japonicus, Amphiprion, 198 
Lophogaster, 577 
Mullus, 509 
Upeneoides, 509 
jeffersoni, Prosciurus, 430 
johannsent, Monohelea, 137, 139, 146 
(fig.), 147 (fig.), 152, 153 
Johannseniella antennalis, 68 
diversa, 76 
nebulosa, 151 
polita, 138 
viridis, 84 
Johannsenomyia polita, 138 
Jonthadocerus, 611 
theryi, 611 
joretiana, Oxytrema, 368 
jourdin, Lutjanus, 194 


125 


kadiakensis, Neomysis, 582 
kansanus, Bolbocerastes imperialis, 106, 
112, 115 (map) 
karafto, Polycelis, 171, 172 
Katerythrops, 581 
sp., 576, 581 
keegani, Geckobia, 10 (fig.), 11 (fig.) 
Kenk, Roman; The freshwater triclads 
(Turbellaria) of Alaska, 163 
Kenkiidae, 566, 568 
Kentrogomphios, 404, 431, 434 
strophensis, 404, 405 (fig.), 406 


_ - _ Mig.), 430 ) y 
kiefferi, Stilobezzia (Stilobezzia), 81 
kincaidi, Boreomysis, 577, 580 
kiushiuana, Upeneoides, 522 
klaas, Lampromorpha, 585, 586, 587, 

590, 592, 593, 594, 595 
knulli, Vasaces, 89, 90, 91 (fig.) 
Kolbeus, 95, 97 

arcuatus, 97, 98 

hornii, 100 

minor, 100 
koltzei, Cephaloon pallens, 161 
koslowi, Polycelis, 172 
kowloonensis, Sermyla, 380 
Kudius, 611 


La Rivers, Ira; Two new naucorid bugs 
of the genus Ambrysus, 1 

labiosa, Paludomus, 369 

Lacais, 611 

glauca, 611 

Lachner, Ernest A.; A revision of the 
goatfish genus Upeneus with 
Pie ae of two new species, 


THE NATIONAL MUSEUM 


VOL. 103 


Lactica, 122, 124, 126, 133, 134 
ferruginea, 123 
intermedia, 123 
scutellaris, 122, 123, 129, 130 
laddi, Fijidoma, 383, 385 
laevigata, Gromphadorhina, 52 
laevissima, Pachychilus, 366 
Lafoéina maxima, 248, 249 
Lagisca extenuata, 222 
extenuata spinulosa, 222 
floccosa, 222 
impatiens, 222, 223 
propinqua, 222 
rarispina, 222 
Lagomorpha, 415, 432 
Lamononia, 611 
monticola, 611 
Lampromorpha, 585, 590, 591, 592, 594 
caprius, 585, 586, 587, 588, 589, 590, 
591, 592, 593 (fig.), 595 
klaas, 585, 586, 587, 590, 592, 593, 
594, 595 
Lampropsephus, 611 
Lanassa, 320, 326 
venusta, 209, 326 
venusta pacifica, 326 
Lanchaeidae, 553 
lanei, Monohelea, 137, 139, 146 (fig.), 
147 (fig.) 
Monohelea (Monohelea), 140, 142 
Lanelater, 608, 611 
schotti, 611 
languarioides, Anischia, 603 
languida, Strabala, 122 
Laphaniella venusta, 326 
lapidum, Glycera, 273 
lapponicus, Ectobius, 41, 45 
laqueata, Oxytrema, 360 
largillierti, Pachychilus, 366 
Sphaeromelania, 365 
Larra, 33 
analis, 32 
Larridae, 32 
Lasiocerus, 611 
lateritia, Tarebia, 378, 379 
laticlavius, Amphiprion, 188, 793 
latidens, KEumys, 411, 412, 413 (fig.) 
latidens, Mesohippus, 430 
latifrons, Dennyus, 533 
latipalpis, Sphaerosyllis, 256, 257 
latrans, Canis, 423 
laurillardi, Tapirus, 479, 481, 485 
Tapirus terrestris, 479 
Lavigeria, 371, 372, 373 
coronata, 372 
grandis, 372 
Lavigeriinae, 358, 369, 371, 373, 386 
Lawrence, R. F.; Two new scale-mite 
parasites of lizards, 9 
lazarus, Scarabaeus, 95, 101 
Leaena, 320, 325 
abranchiata, 209, 325, 326, 331 (fig.) 
abranchiata antarctica, 325, 326 
antarctica, 325 


graffi, 325 
nuda, 326 


INDEX 


lecontei, Bolboceras, 102 
lentigera, Eteone, 235 
Lepidonote cirrata, 220 
scabra, 219 
lepidotus, Elater, 616 
611, 619 
(Aulacon) nobilis, 603 
(Diphyaulon) pyrsolepis, 608 
Le Pinchaque, 471 
Leporidae, 415 
Leptictidae, 403 
leptoceros, Gazella, 424 
Leptochoeridae, 425, 432, 433 
Leptochoerus sp., 425, 430 
Leptomeryx, 427, 432, 433, 434 
annectens, 430 
esuicatus, 426, 427 
evansi, 427 (table) 
mammifer, 427 
iar 426, 427 (table), 
430 
Leptoxis, 361, 363, 364, 369 
praerosa, 361, 362 
lepturides, Cephaloon, 156, 157 (map), 
158, 160 
leschenaulti, Hemidactylus, 12, 15 
Lesnelater, 612 
leucogenys, Pinchacus, 471 
Tapirus, 469, 471, 472, 474, 476, 
477, 479 
leucopeza, Ceratolophus, 138 
Ceratopogon, 138 
Monohelea, 136, 139, 146 (fig.) 
Monohelea (Schizohelea), 138 
Schizohelea, 138 
Leucophaea, 45, 50, 51 
grandis, 51 
maderae, 40, 45, 47 (figs.) 
puerilis, 53 
levis, Harmothoé, 221 
libertina, Semisulcospira, 369 
liebmanni, Pachychilus, 366 
liguriae, Caesaromysides, 577, 581 
limacina, Ophelia, 296 
limbus, Dennyus, 536, 538, 540 (fig.), 542 
limicola, Spio, 284 
Limnetes, 434 
sp., 430 
limnophila, Stilobezzia, 83 
Limnotrochus, 371 
thomsoni, 371 
Lincydrus, 612 
cylindricus, 612 
linearis, Vasaces, 89, 91 (fig.) 
linearis, Elater, 609 
lineata, Pseudosabellides, 318 
lineatus, Elater, 610 
lineolata, Melania, 376 
Hemisinus, 375 
anal aed Aylacostoma (Hemisinus), 
376 


linkoi, Polycelis, 172 
literalis, Pugetomysis, 581 
Lithasia, 363 
geniculata, 363 
obovata, 363 


365272—56——3 


649 


Lithasiopsis, 365 
Lithoglyphus rufofilosus, 387 
litoraurea, Monohelea (Monohelea), 152 
Litorina, 459 
antoni, 458 
cecillei, 456 
malaccana, 456 
nodulosa, 458 
(Tectarius) 
scabra, 568 
thiarella, 451 
tuberculatus, 451 
vilis, 456 
Littoraria, 449, 450 
littorea, Littorina, 450, 460, 462 
Littorina (Littorina), 460 
Littorina, 450, 463 
-capsula habei, 450 
dilatata, 451 
hagruma, 450 
irrorata, 460 
littorea, 450, 460, 462 
(Littorina) littorea, 460 
mauritiana, 450 
multistriata, 450 
(Melarhaphe) neritoides, 450 
nodulosa, 451, 459 
pinctado, 450 
pyramidalis, 451, 456 
rudis, 462 
saxatilis, 462 
seabra, 458 
tuberculata, 451 
ziczac, 450, 453 (fig.) 
(Melarhaphe) ziczac, 450, 452 
Littorinidae, 449, 450, 460, 461, 462 
livens, Blatta, 40, 41 
Ectobius, 39, 40, 47 (figs.) 
livescens, Goniobasis, 361 
livida, Blatta, 40, 41 
Forficula, 41 
lividus, Buffeventius, 604 
Ectobius, 40, 41 
Lizards, Two new scale-mite parasites 


pfeifferianus, 459 


or, 

lobata, Scione, 323 
Lobitarsus, 612 
Lobotarsus, 612 
longa, Apbrodita, 230 

Eteone, 206, 233, 234, 237 (fig.) 

Nereis, 234 
longicauda, Sphaerosyllis, 256, 257 
longiceps, Paedophylax, 259 
longicirris, Paedophylax, 259 
longicornis, Sicardius, 617 
longinaris, Pseudoprotoceras, 430 
longirostris, Lophogaster, 577 
longisetosa, Nephthys, 268 
longisetosus, Autolytus, 242, 247, 250 
Longithorax capensis, 581 
Longiverena, 377 
longosetosa, Nephthys, 268 

Nephthys ciliata, 268 

Nephtys, 207, 267, 268, 269 (fig.) 
samcaaie isa h peaaiag er 242, 247, 

49, 25 


’ 


PROCEEDINGS 


650 


Lophogaster, 577, 578 
affinis, 577 
americanus, 577 
challengeri, 577 
erythraeus, 577 
hawaiensis, 577 
intermedius, 577 
japonicus, 577 
longirostris, 577 
multispinosus, 577 
pacificus, 577 
rotundatus, 577 
schmitti, 577, 578 
spinosus, 577 
subglaber, 577 
subglaber affinis, 577 
subglaber insulare, 577 
typica erythraeus, 577 
typicus, 577 
lourei, Exogone, 259 
lovéni, Melaenis, 205, 206, 214 
Loxorhynchus grandis, 299 
lucidus, Antoligostethus, 603 
Challies, 586, 587, 591, 592, 594, 
59 
lucricrescens, Photuris, 36, 37 
Ludius, 611, 612 
parumcostatus, 611 
lullianus, Cyclopidius, 426 
lumbricalis, Nicomache, 208, 225, 301 
(fig.), 305 
Sabella, 305 
Lumbriconereis minuta, 275, 276 
fragilis, 275 
Lumbricus armiger, 278 
capitatus, 298 
cirratus, 286 
fragilis, 275 
Lumbrinereis fragilis, 275 
Lumbrineris, 275 
fragilis, 207, 275, 277 (fig.) 
Lumbrineridae, 207 (table), 213, 275 
lunulatus, Fulvius, 630 
lutea, Hartomyia, 65, 66 
Stilobezzia, 58, 60, 61, 65, 68 
oe ned, (Neostilobezzia), 65, 72 
g. 
Lutjanus ephippium, 188, 199 
jourdin, 194 
percula, 188, 191 
luzonius, Upeneoides, 519 
Upeneus, 500 (table), 502 (tables), 
503, 504 (table), 505 (tables), 
507, 509, 519, 521 (table), 523 
526 


macfiei, Monohelea, 137, 139, 140, 146 

(fig.), 147 (fig.) 

Monohelea (Monohelea), 143, 144 

macfiei, Stilobezzia, 65 
macrocera, Dido, 600 
macrocerus, Tetralobus, 600, 601 
macropsis, Acanthomysis, 582 
macrostoma, Amphiprion, 201 
macrostomus, Prochilus, 201 
Macrotarsius, 434 


OF THE NATIONAL MUSEUM 


VOL. 103 


maculata, Ampullacera, 385 
Ampullarina, 385 
Fijidoma, 383, 385 
Nicomache, 306 
Pista, 205, 209, 319 (fig.) 323, 325 
Salinator, 385 
Terebella, 323 
Veloplacenta, 385 
maculatus, Vasaces, 89, 91 (fig.), 92 
maculatus, Paludomus, 369 
maculicolle, Cephaloon pallens, 161 
maculipennis, Ceratopogon, 135, 140 
Monohelea, 137, 139, 140 
Monohelea (Monohelea), 140, 142 
Scutops, 555, 556 
madagascariensis, Pachyelater, 612 
maderae, Blatta, 45 
Leucophaea, 40, 45, 47 (figs.) 
Rhyparobia, 45 
Magacnemis, 612 
magnifica, Eusyllis, 205, 207, 260, 261 
Pionosyllis, 261, 262 
magnus, Metacodon, 430 
major, Dennyus, 534, 535, 643 
Eucopia, 579 
Takamatsuia, 543 
malaccana, Litorina, 456 
Nodilittorina, 456 
Maldane, 302, 303 
filifera, 306 
sarsi, 208, 301 (fig.), 303 
sarsi antarctica, 304 
sarsi tropica, 304 
tenuis, 306 
Maldanidae, 208 (table), 213, 301 (figs.), 
302 (key) 
Malekula, 612 
piceus, 612 
Mallaphaga, Studies in Neotropical, 
bird lice of the suborder Am- 
blycera, genus Dennyus Neu- 
mann, 533 
Mallerius, 612 
brasiliensis, 612 
mallochi, Stilobezzia, 65, 66 
Malloea, 600 
malgreni, Prionospio, 207, 282, 283 
g. 
Mammals of northern Colombia, genus 
Tapirus, 465 
memmifer, Leptomeryx, 427 
Manitsha, 434 
Marbenia, 555, 556 
peculiaris, 555, 556 
Margarites, 458 
marginatus, Elater, 607 
mariae, Heliconia, 630 
Marine polychaete worms from Point 
Barrow, Alaska, with additional 
records from the North Atlantic 
and North Pacific, 203 
maritimus, Aplotarsus, 607 
Hypnoidus, 609 
Marshall, N. B., see under Schultz, 
Leonard P. 


INDEX 


marshi, Archaeotherium, 430 
Marsupialia, 402 
mauiensis, Sermyla, 380 
mauritiana, Littorina, 450 
Melarhaphe, 450 

mauritiensis, Amphiprion, 189, 192, 196 
maxima, Lafoéina, 248, 249 
maximus, Abchastus, 601 
maypuri, Tapir, 479, 481, 485 
Mazama, 475 
mecullochi, Amphiprion, 201 
McDermott, Frank A.; Photuris bethan- 

iensis, a new lampyrid firefly, 35 
mediaevus, Apternodus, 403, 430 


medjeorum, Melanoides, 371 melanoleucus, Aeronautes, 538 
Medonia fairmairei, 616 Melanopsidae, 357, 358 
medusa, Polycirrus, 209, 328, 331] Melanopsis, 357, 359 
fig. dufourii, 359 
Megacnemis, 612 melanopus, Amphiprion, 190, 192, 201 
Megalagus, 432, 433, 434 Prochilus, 201 
brachyodon, 416, 430 melanostolus, Amphiprion, 198 
turgidus, 430 Melanotus, 612 
Megapenthes birmanicus, 619 Melanoxanthus, 612 
octomaculatus, 609 rubiginosus, 601 
Megara, 364 virgatus, 609 
alveare, 364 Melanoxocanthus, 612 
megerlei, Brachygonus, 604 melanurus, Amphiprion, 194 
Elater, 604 Melarhaphe, 449, 450, 460, 461 
Melaenis, 214 mauritiana, 450 
lovéni, 205, 206, 214 mentalis, Hoplophoneus, 430 
lovéni gigantea, 214 mercedis, Neomysis, 576, 577, 582 
Melanatractus, 612 meridionalis, Dennyus, 534, 536, 539, 
acutus, 612 540 (fig.), 542 


Melania, 360, 364, 370, 382 


chinensis, 380 
crawshayi, 371 
fasciolata, 380 
ignobilis, 371 
indefinata, 380 
medjeorum, 371 
mweruensis, 371 
sp., 380, 381 
suifuensis, 380 
tornata, 371 


tuberculata, 380, 386 
turriculus, 380, 381 


vainafa, 380 
wagenia, 371 


Nitzschia bruneri, 539 


armigera, 363 Meristhus, 612, 616 


aspirans, 380 
crenocarina, 377 
episcopalis, 382 


glans, 381 Meronychia, 553, 556 
gottschei, 370 Meronychina, 553, 556 
granifera, 379 annulata, 556 
graniperda, 370 Merychippus sp., 423 
hastula, 380 Merycoidodon affinis, 430 


henriettae, 383 
inermis, 381 


oy) 
jacquetiana, 368 Merycoidodontidae, 425, 432, 433 
lineolata, 376 Merycoidodonts sp., 430 
mitra, 380 Mesembria, 601 
nodifila, 370 subtilis, 601 
nodiperda, 370 Mesohippus, 425, 432, 433 
pagodula, 382 celer, 430 


pagodulus, 382 
scalaris, 375 
semigranosa, 379 
siccata, 381 


sulcospira, 381 Mesoreodon chelonyx, 425 
telonaria, 368 messi, Pectocera, 605 
tuberculata, 375, 377 Metacodon, 4381, 433 


variabilis, 382 


quadripunctatus, 617 


scabinula, 616 
scobinula, 612, 616 


culbertsoni, 425 
sp., 425 


651 


hypostylus, 421, 422 (table), 430 


latidens, 430 
montanus, 430 
portentus, 430 


magnus, 430 


Melanian complex, 358 (key) Metactenicerus, 613 


Melanians, The relationships of Old| metallicum, Stenocephaloon, 159 
Meterychrops microphthalma, 580 


and New World, 357 
Melaniidae, 373 


robusta, 580 


Melanoides, 371, 372, 380, 382, 385, | mexianae, Tapirus americanus, 480, 481, 
485 


386 


652 


Microchiroptera, 407 
Microperiscelis, 551, 552, 553, 555, 556 
annulata, 555, 556 
winnertzi, 556 
microphthalma, Meterythrops, 580 
microps, Boreomysis, 576, 57 
Micropternodus, 431, 433 
borealis, 430 
micros, Cryptohypnus, 615 
Microtus, 424 
Midges, American biting, of the heleid 
genus Monohelea, 135 
Biting, of the heleid genus Stil- 
obezzia in North America, 57 
miliaris, Nodilittorina, 450 
milli, Sparus, 198 
mimas, Monohelea (Monohelea), 152 
minimus, Adjidaumo, 430 
minor, Bolbelasmus, 97, 98, 99 (fig.), 
100, 115 (map) 
Bolboceras, 100 
Bradycinetus, 95, 97, 100 
Dennyus, 535, 544 
Ectobius livens, 44 
Kolbeus, 100 
Nicomache, 306 
Nitzschia, 544 
Paradjidaumo, 409, 430 
Perchoerus, 430 
minuta, Aphrodits, 230 
Eulalia, 238 
Eumida, 205, 206, 237 (fig.), 238 
Lumbriconereis, 275, 276 
Oophylax, 256, 257 
Pholoé, 206, 229 (fig.), 230 
minutum, Scalibregma, 293, 294 
minutus, Pseudopterodon, 430 
mirabilis, Cardiohypnus, 604 
mitis, Caenopus, 425, 430 
mitra, Melania, 380 
mobilicornis, Scarabaeus, 101 
Moduliade, 357, 358, 449, 461, 462 
modulus, Modulus, 453 (fig.), 454, 457 
(fig.), 461 
Modulus, 357, 449, 461 
modulus, 453 (fig.), 454, 457 (fig.), 
461 
Mogula sp., 255 
moluccensis, Upeneoides, 514 
Upeneus, 500 (table), 502 (tables), 
504 (table), 505 (tables), 508, 
510, 514 
Upenoides, 514 
monilicornis, Eusyllis, 260 
Syllis, 260 
monofasciatus, Amphiprion, 199 
Monohelea, 135, 736, (key), 139 (table), 
146 (figs.), 147 (figs.) 
American biting midges of the 
heleid genus, 135 
brasiliensis, 137, 135 
(Monohelea) brasiliensis, 145 
guianae, 137, 147 (fig.) 
(Monohelea) guianae, 150 


PROCEEDINGS OF 


THE NATIONAL MUSEUM 


VOL, 103 


hieroglyphica, 135, 136, 137, 139, 
140, 141, 142, 148, 147 (fig.) 

(Monohelea) hieroglyphica, 140, 
144, 145 

hyeroglyphica, 140 

johannseni, 137, 139, 146 (fig.), 147 
(fig.), 152, 153 

lanei, 137, 139, 146 (fig.), 147 (fig.) 

(Monohelea) lanez, 140, 142 

leucopeza, 136, 139, 146 (fig.) 

(Schizohelea) leucopeza, 138 

(Monohelea) litoraurea, 152 

macfiei, 137, 139, 140, 146 (fig.), 
147 (fig.) 

(Monohelea) macfiet, 143, 144 

maculipennis, 137, 139, 140, 146 
(fig.), 147 (fig.) 

(Monohelea) maculipennis, 140, 142 

(Monohelea) mimas, 152 

multilineata, 137, 139, 145, 147 
(fig.), 149, 150 

(Monohelea) multilineata, 149 

nebulosa, 137, 139, 146 (fig.), 147 
(fig.) 151 

(Monohelea) nebulosa, 161, 153 

nigeriae, 150 

ornata, 137, 139, 146 (fig.) 

(Monohelea) ornata, 144 

sp., 135 

stonei, 137, 139, 146 (fig.), 147 (fig.) 

(Monohelea) stonet, 148, 149, 150 

tessellata, 147 (fig.), 150, 152, 153 

(Monohelea) tessellata, 152 

texana, 136, 147 (fig.) 

(Monohelea) texana, 143 
monstrosum, Cavicoxum, 605 
montanus, Glyptosaurus, 401 

Hyanedon, 430 

Mesohippus, 430 

Promerycochoerus, 426 

Stibarus, 430 
monticola, Lamononia, 611 
montrouzieri, Tectaria, 451 
Mopleonus, 613 

candezei, 613 
morgani, Phagocata, 165, 167, 168 
mormon, Ambrysus, 3, 4 
Morrison, J. P. E.; The relationships 

of oS and New World melanians, 
35 
mortoni, Archaeotherium, 430 
moutoniana, Oxytrema, 368 
mucosa, Phyllodoce, 236 
Mudalia, 361 

carinata, 361 

trilineata, 361 
Mulloides pinnivittatus, 513 
Mullus, 528 

auratus, 528, 529 

bandi, 516 

bensasi, 509 

dubius, 498, 515 

japonicus, 509 

subvittatus, 513 

vittatus, 506, 516 


INDEX 


653 


multilineata, Monohelea, 137, 139, 145, | Neolacon, 613 


147 (fig.), 149, 150 
Monohelea (Monohelea), 149 
Oxytrema virginica, 360 
Palpomyia, 149 
multipapillata, Castalia, 239 
multisetosa, Harmothoé, 223 
multispinosus, Lophogaster, 577 
multistriata, Littorina, 450 
Mumetopia, 557, 558 
muricatus, Tectarius, 449, 450, 452, 453 
(fig.), 454, 455 (fig.), 457 (fig.), 
459, 460, 462 
murinus, Elater, 602 
Mustelinae, 434 
Mutiloptera, 557 
mweruensis, Melanoides, 371 
Mycterus, 156, 158 
Myodris, 552, 553, 555, 556 
annulata, 556 
Myrsidea, 533 
A supplement to W. M. Tattersall’s 
review of, 575 
Mysidae, 582 
Mysidella americana, 576, 582 
Mysidellinae, 576, 582 
Mysini, 576 
Mysis oculata, 576, 581 
Mysta, 233 
barbata, 233 
Mystides, 231, 232 
borealis, 206, 232, 237 (fig.) 
notialis, 232 
viridis, 232 
Myxicola, 334, 340 
infundibulum, 209, 340, 341 (fig.) 
steenstrupii, 340 


naidina, Exogone, 207, 243 (fig.), 258 
nana, Glycera, 273 
nanus, Perchoerus, 430 
nassa, Nassopsis, 372 
Nassopsis, 373 
nassa, 372 
Naucorid bugs of the genus Ambrysus, 
Two new, 1 
Naucoridae, 1 
Nauphoeta, 48, 51 
cinerea, 40, 46, 51 
nausicaae, Dendrocoelum, 179 
nearcticum, Rhopalosoma, 19, 20, 24, 32 
nebulosa, Ceratopogon, 135 
Cyamops, 558 
Hartomyia, 151 
nebulosa, Johannseniella, 151 
Monohelea, 137, 139, 146 (fig.), 
147 (fig.), 151 
Monohelea (Monohelea), 151, 153 
Periscelis, 553, 556 
nebulosus, Ceratolophus, 151 
Ceratopogon, 151 
neglecta, Potamilla, 209, 333 (fig.), 335 
Sabella, 335 
neglectus, Pseudocylindrodon, 430 
Neoamphitrite robusta, 322 


formosanus, 613 
Neomysis franciscorum, 582 

kadiakensis, 582 

mercedis, 576, 577, 582 

rayli, 582 
neonigripes, Nereis, 264 

Nereis pelagica, 264, 265 
Neonomopleus, 613 

strictus, 613 
Neoscutops, 555, 556 

rotundipennis, 555, 556 
Neostilobezzia, 63 
Neotropical Miridae, LXIV, New bugs 

of the subfamily Cylapinae 
(Hemiptera), 621-632 

nephrophthalma, Acanthomysis, 582 
Nephthys brachycephala, 271, 272 

caeca, 270 

ciliata, 268, 270 

ciliata longosetosa, 268 

discors, 270 

coeca, 268 

longisetosa, 268 

longosetosa, 268 

paradoxa, 271 

phyllobranchia, 271 
Ne 207 (table), 212, 266, 269 


gs. 
Nephtys, 267 (key) 
ciliata, 207, 267, 269 (fig.), 270 
discors, 207, 267, 269 (fig.), 270 
longosetosa, 207, 267, 268, 269 


g. 
paradoxa, 207, 267, 269 (fig.), 271 
phyllobranchia, 271, 272 
rickettsi, 270, 271 

Nepthys ciliata, 270 
hudsonica, 270 

pais: 207 (table), 212, 263, 269 


(figs.) 

Nereis, 263, 264 (key) 

aphroditoides, 239 

arctica, 264 

armillaris, 254 

caeca, 270 

ciliata, 268, 270 

filicornis, 284 

flava, 235 

longa, 234 

neonigripes, 264 

pelagica, 207, 264, 269 (fig.) 

(Nereis) pelagica, 264 

pelagica neonigripes, 264, 265 

pelagica occidentalis, 264, 265 

prismatica, 247, 249 

procera, 266 

zonata, 207, 265, 269 (fig.) 

(Nereis) zonata, 264, 265 

zonata persica, 266 
Nerinides, 280, 285 

sp., 205, 207, 283 (fig.), 285 
neritiformis, Tylomelania, 381 
neritinoides, Stanleya, 388 
neritoides, Littorina (Melarhaphe), 450 


654 


Nicolea, 320, 322 
arctica, 322 
simplex, 323 
venustula, 209, 319 (fig.), 322 
zostericola, 322 
Nicomache, 303, 304, 305 (key) 
carinata, 305 
lumbricalis, 208, 225, 301 (fig.), 306 
lumbricalis borealis, 305 
maculata, 306 
minor, 306 
personata, 208, 301 (fig.), 305, 306 
niger, Pleonomus, 616 
nigeriae, Monohelea, 150 
nigra, Polycelis, 172 
nigrata, Pachychilus, 366 
nigriceps, Strabala, 122 
Nimravinae, 434 
Nina, 458 
Nitzschia bruneri, 538 
bruneri meridionalis, 539 
dubius, 537 
minor, 544 
pulicaris tibialis, 538 
tibialis, 538 
nivea, Phagocata, 164, 167 (fig.), 179, 
180, 182 
nobilis, Elater, 601 
Lepidotus (Aulacon), 603 
nodifila, Melania, 370 
Oxytrema, 361, 368 
Nodilittorina, 449, 450, 454, 455 (fig.), 
458, 459, 460, 463 
malaccana, 456 
miliaris, 450 
picta, 450 
pyramidalis, 456, 460 
subnodosa, 456 


tuberculata, 449, 450, 451, 453 
(fig.), 454, 455 (fig.), 457 (fig.), 
459, 460, 462 

vilis, 456 


nodiperda, Melania, 370 
nodosa, Eunoa, 217 
Eunoé, 206, 216, 217, 219, 229 (fig.) 
Harmothoé, 217, 219 
Polynoé, 217, 219 
nodulosa, Litorina, 458 
Littorina, 451, 459 
nodulosus, Echininus, 449, 451, 452, 
453 (fig.), 454, 455 (fig.), 457 
(fig.), 458 
Tectarius, 462 
Trochus, 451, 456 
Turbo, 451 
Nomopleus, 613 
nordmanni, Goniada, 274 
North American triclad Turbellaria, 
XIII; three new cave planarians, 
563 
Northern Colombia, Mammals of, genus 
Tapirus, 465 
Nothocyon geismarianus, 419 
notialis, Mystides, 232 
Notiphila, 553, 556 
annulata, 552 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL, 103 


nubilus, Canis, 423 
nuda, Leaena, 326 
Nullarborica, 613 
concinna, 613 
Nychia cirrosa, 226 
Nyctitheriidae, 404 
Nyctor, 613 
expallidus, 613 


obesus, Elater, 608 
obovata, Lithasia, 363 
obscura, Pacificola, 613 
obscuripes, Elater, 605 
obscurus, Elater, 606 
occidentalis, Periscelis, 552, 553, 554, 
555, 556 
occidentalis, Nereis pelagica, 264, 265 
ocellaris, Amphiprion, 194 
ocellata, Wetmorella, 440, 442, 443, 444, 
446, 447 
Ochotoninae, 434 
ochraceus, Ceratopogon, 63 
octomaculatus, Megapenthes, 609 
oculata, Mysis, 576, 581 
oculatus, Vannius, 626 (fig.), 628 
oculi-marginata, Polycelis, 172, 
Odocoileus hemionus, 423 
virginianus, 423 
Odontaeus, 96, 101 
Odontocardus, 613 
Oedemeridae, 158 
oerstedi, Chaetosyllis, 253 
Eunoé, 206, 216, 218, 219, 229 
(fig.) 
Syllis, 253 
Ogmophis arenarum, 402 
o’harrai, Hoplophoneus, 430 
olequaensis, Ambloxus, 360 
oligospilus, Upeneus, 500 (table), 502 
(tables), 503, 504 (table), 505 
(tables), 509, 525, 526 (table), 
527 (table) 
onager, Equus, 424 
Oophylax minuta, 256, 257 
Ophelia limacina, 296 
eee a (table), 213, 295 (key), 


g. 
Ophelina groenlandica, 295, 296 
Opomyzidae, 557 
Orbiniidae, 207 (table), 213, 278, 283 


(figs.) 
pretilies Asabellides, 318 
ornata, Monohelea, 137, 139, 146 (fig.) 
Monohelea (Monohelea, 144 
ornamentatus, Idanthyrsus, 308 
ornata, Proceraea (Stephanosyllis, 
Stephanosyllis, 246 
ornatifrons, Fulvius, 626 (fig.), 631 
ornatus, Anchastelater, 602 
Autolytus, 244 
Diploconus, 616 
Tremarctos, 475 
Orocharis gryllodes, 24 
saltator, 22, 23, 24 
sp., 22, 23, 24, 25, 32 
osborni, Trigonias, 425, 430 


182 


246 


INDEX 


osculata, Goniobasis, 361 
ovalis, Elater, 602 
Ovalpalpus, 613 
Ovipalpus, 613 
ovoideum, Gyrotoma, 361 
Oxymelania, 365 
schiedianus, 365 
Oxytrema, 359, 364, 367, 370, 371, 382 
bailleti, 368 
bulbosa, 361 
canaliculatum undulatum, 360 
catenaria, 360 
comalensis, 360 
deshayesiana, 361 
jacquetiana, 368 
joretiana, 368 
laqueata, 360 
moutoniana, 368 
nodifila, 361, 368 
peregrinorum, 368 
(Strephobasis) plena, 361 
plicifera, 360 
semicarinata, 367 
silicula, 360 
symmetrica, 360 
telonaria, 368 
toucheana, 368 
virginica, 364 
virginica multilineata, 360 


Pachychilus, 360, 364, 366, 367, 370 
corvinus, 365, 366 
cumingii, 366 
(Pilsbrychilus) dalli, 367 
drakei, 360 
fuentesi, 366 
glaphyrus, 365 
graphium, 366 
laevissima, 366 
largillierti, 366 
liebmanni, 366 
nigrata, 366 
pilsbryi, 365 
(Oxymelania) pilsbryi, 376 
violaceus, 366 
walli, 367 
Pachycynodon, 421 
Pachyelater, 612 
madagascariensis, 612 
pacifica, Euchaetomeropsis, 577, 580 
Lanassa venusta, 326 
Spio filicornis, 284 
Pacificola, 613 
obscura, 613 
pacificum, Cephaloon, 157 (map), 159, 160 
pen Lophogaster, 577 
aedophylax dispar, 259 
longiceps, 259 
longicirris, 259 
pagodula, Protia, 382 
Melania, 382 
pagodulus, Melania, 382 
Paleocastor gradatus, 408 
peninsulatus, 408 
Paleogale, 432, 433 
inflex, 430 


Paleolagus, 432, 433, 434 
burkei, 416 
haydeni, 415 
intermedius, 415 
temnodon, 414, 430 


655 


ae aE alee 157 (map), 158, 160, 


Sanateen ies Hartomyia, 78 
Stilobezzia, 59, 61, 78 
Stilobezzia (Stillobezzia), 78 

pallidulus, Elater, 602 

Palpomyia dorsofasciata, 62 
multilineata, 149 

Paludominae, 371 

Paludomus, 369, 383 
labiosa, 369 
maculatus, 369 
tanschaurica, 369 

Panamenia, 555, 556 
chapmani, 555 

Panchlora, 45, 51 

Pantolamprus sulcicollis, 618 

panzeri, Ectobius, 44 

papuensis, Amphiprion, 199 
Hortizoteichos, 610 

Parabezzia, 61 
elegantula, 62 
petiolata, 61 
poikiloptera, 62, 83 

Paracapiophonus, 614 

Paracardiophorus, 614 

Paracrostoma, 383 
hugeli, 383 

Paradaxon, 614 

Paradoxon, 614 

Paradima, 614 
tattakensis, 614 

Paradjidaumo, 433 
minor, 409, 430 
spokanensis, 409 
trilophus, 409 

paradoxa, Nephthys, 271 


Nephtys, 207, 267, 269 (fig.), 271 


Paragonimiasis, 3 
Paralithodes camtschatica, 224 
Paramelania, 371, 372, 373 
damoni, 371 
Paramelaniinae, 373 
Paranaitis wahlbergi, 236 
Parapomachilius, 601 
Parascaptor, 407 
Parastirus, 614 
Parathous, 614 
sanguineus, 614 
Parcoblatta, 43, 45, 50 
Parictis, 421, 432, 433 
bathygenus, 421 
dakotensis, 420, 430 
Paroedostethus, 614 
relictus, 614 
parumcostatus, Ipostirus, 611 
Ludius, 611 
Parupeneus, 515 
parviceps, Ambrysus, 6 
Parvistoma, 614 


656 


parvulus, Bolborhombus, 99 (fig.), 115 
(map), 117, 118, 119 
parvus, Upeneoides, 528 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 108 


wheeleri, 552, 653, 556 
winnertzi, 556 
Perissodactyla, 421, 432 


Upeneus, 499, 500 (table), 501, 502] Peritropis, 624 


(tables), 503, 504 (table), 505 
(tables), 508, 528 

paterculus, Hesperocyon, 4/6, 417 (fig.), 
430 


Patricia, 614 
austratica, 614 

paucicostata, Edgaria, 372 

paulistensis, Stilobezzia, 71 

paululus, Hypisodus, 430 

Peasiella, 462 

Pectinaria, 311 
(Cistenides), 311 (key) 
brevicoma, 312 
(Cistenides) brevicoma, 312 
eranulata, 208, 313 (fig.) 
(Cistenides) granulata, 311, 312 
hyperborea, 208, 313 (fig.) 
(Cistenides) hyperborea, 311, 314 
sp., 307 

Pectinariidae, 208 (table), 212, 310, 313 

s 


figs. 
pectinicornis, Elater, 606, 607 
Pectocera, 614 
messi, 605 
Pectoerra, 614 
Pectora, 614 
peculiaris, Marbenia, 555, 556 
pelagica, Chaetura, 537 
Nereis, 207, 264, 269 (fig.) 
Nereis (Nereis), 264 
pellucida, Planaria, 167 
Peltosaurus sp., 401 
Penecurva, 168 
Pengamethes, 614 
fulvicollis, 614 
peninsularis, Bolbocerastes, 99 (fig.), 
105, 112, 114 (map), 116 
Bolboceras serratus, 116 
Bradycinetus serratus, 116 
peninsulatus, Paleocastor, 408 
Peratherium fugax, 402 
titanelix, 402, 430 
Perea polymna, 194 
Perchoerus, 433 
minor, 430 
nanus, 430 
percula, Actinicola, 191 
Amphiprion, 188, 197, 192, 193 
Lutjanus, 188, 191 
Prochilus, 191 
pereerinorum, Oxytrema, 368 
perideraion, Amphiprion, 188, 191, 192 
Prochilus, 191 
Periplaneta, 46, 50, 53 
Periscelidae, 551, 652, 555 (key), 556 
(list), 557, 560 
Periscelis, 551, 552 (key), 555, 556 
annulata, 552, 653, 556 
annulipes, 556 
nebulosa, 553, 556 
occidentalis, 552, 553, 564, 555, 556 


Peritropoides, 622 
annulatus, 624, 625, 628 
quadrinotatus, 625, 626 (fig.), 627 
(fig.), 629 (fig.) 
Peromyscus, 410 
persica, Nereis zonata, 266 
personata, Nicomache, 208, 301 (fig.), 
305, 306 
perspicillaris, Eetobius, 41 
peruvianus, Tapirus (terrestris), 479, 


’ 
Petaloproctus, 302, 306 
filifer, 306, 307 
tenuis, 208, 301 (fig.), 306 
tenuis borealis, 306, 307 
petiolata, Parabezzia, 61 
Pettibone, Marian H.; Marine poly- 
chaete worms from Point Barrow, 
Alaska, with additional records 
from the North Atlantic and 
North Pacific, 203 
pfeifferianus, Litorina (Tectarius), 459 
Phagocata, 164, 166, 167, 168, 566 
albissima, 165, 167 
cavernicola, 563, 565 (fig.) 
coarctata, 167 
morgani, 165, 167, 168 
nivea, 164, 167 (fig.), 179, 180, 182 
subterranea, 566 
vitta, 165, 167 
Phalerebus, 188, 190 
Pharotarsus, 614 
phaseopygus, Chaetura  griseiventris, 
539 


Phedomenus flavangulus, 608 

Pherhimius, 615 

Phibisa, 615 

philippina, Wetmorella, 439, 440, 446, 
447 


Wetmorella philippina, 440, 441 
(fig.), 442, 443, 445 
philippinensis, Geckobia, 12, 13 (fig.), 
14 (fig.) 
philippinus, Upeneoides, 516 
phillipsi, Upeneus, 498, 528, 529 
Philodendron sp., 631 
phoenicobia, Tachnornis phoenicobia, 
543 


Pholoé, 228, 230 

minuta, 206, 229 (fig.), 230 

tuberculata, 230 
Phorocardius, 615 

solitarius, 608 
Phorotarsus, 614 
Phorticoides, 556 

flinti, 553, 556 
phosphorea, Eusyllis, 260, 261 
Photinus consanguineus, 36 

pyralis, 35, 36 
Photuris, 35 

bethaniensis, 36 


INDEX 


bethaniensis, a new Lampyrid fire- 


fly, 25 
fairchildi, 37 
hebes, 36 
lucricrescens, 36, 37 
versicolor, 35, 36 
phyllobranchia, Nephthys, 271 
Nephtys, 271, 272 
Phyllodoce, 236 
citrina, 236 
groenlandica, 206, 236, 237 (fig.) 
(Anaitides) groenlandica, 236 
mucosa, 236 
Phyllodocidae, 206 (table), 
(key), 237 (figs.) 
Piaya, 586 
piceus, Malekula, 612 
picta, Hartomyia, 75 
Nodilittorina, 450 
Stephanosyllis, 246 
picticolle, Cephaloon pallens, 161 
pictus, Ceratopogon, 75 
piegeti, Dennyus, 533 
Piezophyllus, 600, 601, 606 
spencei, 601, 606 
pilosa, Brada, 290, 291 
Pilsbrychilus, 366 
pilsbryi, Pachychilus, 365 
Pachychilus (Oxymelania), 376 
ibe sea Tapirus, 471, 480 
inchacus, 469, 471 
aenigmaticus, 471 
leucogenys, 471 
pinchaque, 471 
terrestris, 471 
villosus, 471 
pinchaque, Pinchacus, 471 
Tapir, 471 
Tapirus, 465, 466, 467, 468, 469, 
471, 478, 479, 482 (fig.), 483 
(fig.), 484, 487 (fig.), 489, 493 
(map), 495 (map) 
pinctado, Littorina, 450 
pinnivittatus, Mulloides, 513 
Pionosyllis, 242, 262 
compacta, 205, 207, 243 (fig.), 262 
magnifica, 261, 262 
pirtformis, Dendrocoelopsis, 174, 177 
(fig.), 178, 180 
Pista, 320, 323 
flexuosa, 226 
groenlandica, 324 
cae cia 205, 209, 319 (fig.), 323, 
5 
Placonides, 615 
Planaria pellucida, 167 
Planariidae, 164, 563 
Planaxidae, 357, 358, 387 
Platycrepidius, 618 
eet Oxytrema (Strephobasis), 361 
leonomus argentatus, 613 
niger, 616 
Plesictis, 482, 433 
priscus, 430 
Pleurocera, 359, 360, 362, 363 
acuta, 360 


212, 231 


657 


(Oecytrema) acuta, 260 
alveare, 364 
(Ellipstoma) gibbosa, 363 
verucosa, 362 
zonalis, 363 
Pleuroceridae, 357, 358, 359, 363, 365, 
867, 369, 370, 374, 375, 376, 377 
pare iene 358, 359, 364, 367, 369, 
3 
Pleurocerus (Oxytrema) acutus, 360 
pliacus, Ischyromys, 408, 430 
plicifera, Oxytrema, 360 
Ploconides, 615 
Plotia, 379 
Plotiopsis, 378 
balonnensis, 378 
pluribranchiata, Stylaroides, 290, 291 
Podocera, 556, 560 
ramifera, 560 
Poébrotherium, 433 
eximium, 426, 430 
poeyi, Phopalosoma, 19 
poikiloptera, Parabezzia, 62, 83 
polita, Allohelea, 138 
Johannseniella, 138 
Johannsenomyia, 138 
Schizohelea, 138 
Sphaeromias, 138 
politus, Ceratolophus, 138 
Ceratopogon, 138 
Polybostrichus, 241, 247, 248, 251 (fig.) 
longosetosus, 242, 247, 249, 250 
Polycelidia, 171 
Polycelis, 163, 168, 169, 171, 172, 173, 
182, 183 (map), 184 
auriculata, 171, 172 
borealis, 168, 171 (fig.), 172, 182 
coronata, 171, 172 
eburnea, 172 
elongata, 172 
eudendrocoeloides, 172 
eurantron, 172 
felina, 172 
karafto, 171, 172 
koslowi, 172 
linkoi, 172 
nigra, 172 
oculi-marginata, 172, 182 
polyopis, 172 
receptaculosa, 172 
relicta, 172 
sabussowi, 172 
sapporo, 172 
schmidti, 171, 172 
tenuis, 172 
tibetica, 172 
Polychaeta, 211 (key) 
Polycirrus, 320, 328 
medusa, 209, 328, 331 (fig.) 
sp., 328 
Polydora, 280, 281 
brachycephala, 280, 281 
caulleryi, 207, 280, 281, 283 (fig.) 
polylepis, Prochilus, 200 
polymna, Anthias, 198 
Perca, 194 


658 


polymnus, Amphiprion, 189, 193, 194, 
197, 198, 200 
Anthias, 191 
Prochilus, 198 
Polynoé arctica, 217 
cirrata, 220 
extenuata, 222 
floccosa, 222 
foraminifera, 217 
gracilis, 225 
islandica, 217, 218 
nodosa, 217, 219 
rarispina, 222 
sarsi, 215 
scabra, 217, 218, 219 
semisculpta, 222 
semisculptus, 222 
spinulosa, 217 
Polynoidae, 206 (table), 212, 213, 214 
(key), 229 (figs.) 
polyopis, Polycelis, 172 
Polyprotodontia, 402 
portentus, Mesohippus, 430 
Potadoma, 370 
freethii, 370 
Potadominae, 371 
Potamanax, 365, 376 
Potamididae, 449 
Potamilla, 334, 335 
acuminata, 336 
neglecta, 209, 333 (fig.), 385 
reniformis, 209, 333 (fig.), 335, 336 
torelli, 336 
Practapyrus, 615 
descarpentriesi, 615 
praerosa, Leptoxis, 361, 362 
praetermissa, Clymene (Praxillella), 303 
Praxilla, 303 
Praxillella, 208, 301 (fig.), 303 
Praxilla praetermissa, 303 
Praxillella, 302, 303 
praetermissa, 208, 301 (fig.), 303 
prezevalskii, Equus, 424 
prima, Diopsosoma, 556 
Prionospio, 280, 282 
malmgreni, 207, 282, 283 (fig.) 
tenuis, 282 
priscus, Plesictis, 430 
prismatica, Nereis, 247, 249 
prismaticus, Autolytus, 207, 242, 244, 
245, 246, 247, 249, 251 (fig.) 
Pristilophus, 615 
sericans, 616 
Pristolophus, 615 
Probezzia elegantula, 62 
procera, Nereis, 266 
Proceraea gracilis, 250 
(Stephanosyllis) ornata, 246 
Prochilos ephippium, 199 
Prochilus, 188 
akallopisos, 188, 190 
bifasciatus, 193, 194 
macrostomus, 201 
melanopus, 201 
percula, 191 
perideraion, 191 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL, 103 


polymnus, 198 
polypepis, 200 
Proclea, 320, 326 
graffi, 325 
graffii 209, 325 
Prodrasterius, 615 
Progne chalybea chalybea, 549 
Prolacon, 615 
allaudi, 615 
aie ep A 244, 245, 247, 249, 


Promerycochoerus montanus, 426 
Proneomysis wailesi, 582 
propinqua, Harmothoé, 222 
Harmothoé rarispina, 222 
Lagisca, 222 
Proquasimus, 615 
Prosciurus, 431, 433 
jeffersoni, 430 
relictus, 408 
vetustus, 430 
Prosternon, 600 
Prosynthetoceros, 428 
Protemnocyon, 420 
pruinosa, Stilobezzia, 59, 60, 61, 79 
Stilobezzia (Stilobezzia), 73 (fig.), 
79 
Psammate aphroditoides, 239 
Psephus cyaneus, 611 
Pseudathous, 615 
Pseudocylindrodon, 432, 433 
neglectus, 430 
Pseudoderomecus, 616 
Pseudo-Elater, 616 
pseudomacropsis, Acanthomysis, 582 
Pseudomma truncatum, 581 
Pseudonomopleus, 616 
Pseudoplotia, 378 
scabra, 378 
Pseudopotamilla intermedia, 337 
reniformis, 337 
Pseudopristilophus, 616 
Pseudoprotoceras, 433, 434 
longinaris, 430 
Pseudopterodon, 434 
minutus, 430 
Pseudosabellides, 318 
lineata, 318 
Pseudostirus, 616 
Pterygosomidae, 10 
pudicus, Ambrysus, 6 
puerilis, Leucophea, 53 
puertoricensis, Strabala ambulans, 125 
(fig.), 127, 132 
Pugetomysis literalis, 581 
pulchripennis, Sphaeromyas, 137 
Pulchronotus, 616 
puncticollis, Ambrysus, 6 
punctipes, Stilobezzia, 58, 60, 61 
Stilobezzia (Stilobezzia), "72 (fig.), 79 
punctulata, Stilobezzia (Stilobezzia), 81 
pupa, Travisia, 296 
pupieri, Ctenicera, 615 
purpureua, Elater, 614 
Pycnoscelus, 45, 51 
surinamensis, 45, 51 


INDEX 


pyralis, Photinus, 35, 36 
pyramidalis, Littorina, 451, 456 
Nodilittorina, 456, 460 
pyrsolepis, Adelocera, 608 
Lepidotus (Diphyaulon), 608 
Pythidae, 158 


quadrangularis, Spirorbis, 343 

quadratus, Hapithus agitator, 23, 24 

quadridens, Bolboceras, 101 

Scarabaeus, 101 

quadrinotatus, Peritropoides, 625, 626 
(fig.), 627 (fig.), 629 (fig.) 

quadripunctatus, Meristhus, 617 

quadristillatus, Fulvius, 627 (fig.), 631 

quadrivittatus, Betarmon, 611 

Quasimus setosus, 606 

quaternaria, Syllis, 253, 254 


rabelloi, Stilobezzia, 59, 60, 61, 81 
Stilobezzia (Stilobezzia), 72 (fig.), 81 
radiata, Amphitrite, 321 
Radina, 380 
ramifera, Podocera, 560 
Rangifer arcticus, 424 
caribou, 424 
Raphaea, 616 
vitida, 616 
rarispina, Harmothoé, 222 
Lagisca, 222 
Polynoé, 222 
Rastrocephalus, 616 
rayii, Neomysis, 582 
receptaculosa, Polycelis, 172 
rectangularis, Colaulon, 606 
reedi, Eotylopus, 430 
regalis, Bolbocerastes, 99 (fig.), 
106, 109, 112, 115 (map) 
relicta, Polycelis, 172 
relictus, Paroedostethus, 614 
Prosciurus, 408 
reniformis, Potamilla, 209, 333 (fig.), 
335, 336 
Pseudopotamilla, 337 
Samella, 336 
Reptilia, 401 
revelieri, Athous, 603 
Review of the Atlantic periwinkles, 
Nodilittorina, Echininus, and 
Tectarius, 449 
rez, Bradycinetulus, 99 (fig.), 102, 103 
114 (map) 
Rhaciaspis, 616 
Rhinoceretidae, 425, 432, 433 
Rhinochoerus, 477 
Rhino(choerus) villosus, 471 
Rhopalosoma, 19, 20, 21, 22, 23, 24, 25, 
27 (figs.), 29 (figs.), 31, 32, 33 
aenigmaticum, 19 
nearcticum, 19, 20, 24, 32 
Notes on the biology and immature 
stages of a cricket parasite of 
the genus, 19 
poeyi, 19 
Rhopalosomatidae, 19 
Rhopalosomidae, 19 


105, 


659 


Rhyparobia, 45 
maderae, 45 
richmondi, Chaerura, 537 
Dennyus, 533, 535, 537 
rickettsi, Nephtys, 270, 271 
riparius, Elater, 606, 610 
Risella, 462 
Risellidae, 462 
Riselloidea, 462 
Risellopsis, 462 
Rismethus, 616 
Rjabuschinskya, 172 
robusta, Amphitrite, 322 
Meterythrops, 580 
Neoamphitrite, 322 
robustus, Cirratulus, 286 
Hoplophoneus, 430 
Rodentia, 408, 431 
Roggeveenia, 616 
buxtoni, 616 
rohan-chaboti, Ameidioides, 602 
rosenbergi, Amphiprion, 191 
Rostricephalus, 616, 617 
vitalisi, 617 
rotunda, Strabala, 125 (fig.), 127, 130, 
133 


rotundatus, Lophogaster, 577 
rotundipennis, Neoscutops, 555, 556 
rotundocapitis, Dennyus, 535, 541 (fig.), 
548, 549 
roulini, Tapirus, 471, 473, 476, 480 
roulinii, Tapirus, 466, 471, 472, 476, 479 
rubiginosus, Melanoxanthus, 601 
rubrocinectus, Amphiprion, 199 
rubrovittatus, Vannius, 629 
rudis, Littorina, 462 
rufa, Disonycha, 122 
Haltica, 121, 122, 123, 127 
Strabala, 125 (fig.), 126, 127, 128, 
129 
ruficollis, Agonischius, 605 
rufofilosa, Tanganyicia, 387 
rufofilosus, Lithoglyphus, 387 
rufus, Tapirus, 479, 481, 485 
ruginosum, Aylacostoma, 377 
rugosa, Brada, 290, 291, 292 
Trophonia, 290 
rippeli, Amphiprion, 199 
Rygodonus, 617 
Rymcobites, 617 
singularis, 617 


sabatyra, Tapyra, 479, 481, 485 
sabella, Fabricia, 299 
Sabella, 334 
analis, 339 
crassicornis, 209, 333 (fig.), 334 
fabricii, 334 
granulata, 312 
lumbricalis, 305 
neglecta, 335 
reniformis, 336 
spetsbergensis, 334 
spitzbergensis, 334 
Sabellariidae, 208 (table), 


212, 301 
(figs.), 307 


660 


Sabellidae, 209 (table), 211, 332, 333 
(figs.), 334 (key), 341 (fig.) 
Sabellides sibirica, 318 
sabussowi, Polycelis, 172 
Sacconereis, 241, 246, 248, 250, 251 (fig.) 
Saccopsis terebellidis, 330 
sakurae, Cephaloon, 161 
Salinator maculata, 385 
saltator, Orocharis, 22, 23, 24 
sanguineus, Parathous, 614 
sanguinipennis, Georgicus, 609 
Saphromyzidae, 553 
sapporo, Polycelis, 172 
sarsi, Antinoé, 205, 206, 215 
Antinoélla, 215 
Eteone, 235 
Harmothoé, 215 
Harmothoé (Antinélla), 215 
Maldane, 208, 301 (fig.), 303 
Polynoé, 215 
Sauria, 401 
saxatilis, Aeronautes s., 537, 538 
Littorina, 462 
scabinula, Meristhus, 616 
seabra, Aphrodita, 217 
Lepidonote, 219 
Litorina, 458 
Littorina, 458 
Polynoé, 217, 218, 219 
Pseudoplotia, 378 
scalare, Aylacostoma (Aylacostoma), 
6 


scalaris, Melania, 375 
Seale-mite parasites of lizards, Two 
new, § 
Scalibregma, 293 
brevicauda, 293, 294 
inflatum, 208, 293, 297 (fig.) 
mintum, 293, 294 
Scealibregmidae, 208 (table), 213, 293, 
7 (figs.) 
Scalopinae, 403 
Sealopus aquaticus, 404 
Scapatochirus, 407 
Scarabaeus cephus, 101 
cyclops, 101 
farctus, 101 
ferrugineus, 101, 102 
gallicus, 97 
lazarus, 95, 101 
mobilicornis, 101 
mobilicornis testaceus, 101 
quadridens, 101 
Seeloporus, 15 
undulatus, 9, 15 
schaefferi, Bolboceras, 117, 119 
Bolborhombus, 99 (fig.), 114 (map), 
117, 119 
schiedianus, Oxymelania, 365 
Schizohelea, 137 
copiosa, 138 
leucopeza, 138 
polita, 138 
schmidti, Polycelis, 171, 172 
schmitti, Lophogaster, 577, 578 
schotti, Agryphus, 611 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 103 


Lanelater, 611 
schusteri, Cardiophorus, 611 
scobinula, Meristhus, 612, 616 
Scoliocerus, 603 
scotti, Archaeotherium, 430 
Schultz, Leonard P.; Review of the 
Indo-Pacific anemone fishes, 
genus Amphiprion, with descrip- 
tion of two new species, 187 
And Marshall, N. B.; A review of 
the labrid fish genus Wet- 
morella with descriptions of new 
forms from the tropical Indo- 
Pacific, 439 
Sciaena unimaculata, 194 
Scione lobata, 323 
Sciuravidae, 408 
Scoloplos, 278 
armiger, 207, 278, 283 (fig.) 
sculpta, Acanthomysis, 582 
sculpticauda, Eucopia, 578 
scutata, Sternaspis, 208, 309, 313 (fig.) 
Thalassema, 309 
scutellaris, Altica, 121 
Haltica, 123 
Lactica, 122, 123, 129, 130 
Strabala, 122, 123 
scutellata, Strabala, 121 
Seutops, 555, 556 
chapmani, 556 
fascipennis, 555, 556 
maculipennis, 555, 556 
sebae, Amphiprion, 189, 192, 197 
Seidlia, 172 
Selatosomus guttatus, 610 
semicarinata, Oxytrema, 367 
semigranosa, Melania, 379 
Tarebia, 379 
Semiotinus, 617 
semisculpta, Polynoé, 222 
semisculptus, Polynoé, 222 
Semisinus, 375 
Semisulcospira, 369 
libertina, 369 
Semisulcospirinae, 369 
sequax, Ceratopogon, 135 
sericans, Pristilophus, 616 
Sermyla, 380 
kowloonensis, 380 
mauiensis, 380 
tornatella, 380 
Serpentes, 402 
Serpula granulata, 343 
spirillum, 344 
Benin 209 (table), 211, 341 (figs.). 
42 


serrata, Inusitatomysis, 581 
serraticornis, Dicronychus, 609 
serratus, Amechanus, 113 
Athyreus, 113 
Athyreus (Bradicinetus), 113 
Bolboceras (Amechamus), 113 
Bolbocerastes, 99 (fig.), 106, 107, 
108, 109, 112, 173, 115 (map) 
Bradycinetulus, 113 
Bradycinetus, 113 


INDEX 


Serropalpidae, 159 
Setaera. 379 
setosa, Brada, 290, 291 
Chaetozone, 207, 287, 297 (fig.) 
Glycera, 272 
Spio, 284 
Thiara, 379 
setosus, Quasimus, 606 
Sheppardiconcha, 367 
sibirica, Asabellides, 208, 318, 319 (fig.) 
Sabellides, 318 
Sicardius, 617 
longicornis, 617 
siccata, Melania, 381 
siculus, Ectobius, 41 
Sigalionidae, 206 (table), 211, 228 
signoreti, Ambrysus, 6 
silicula, Oxytrema, 360 
* si marmmanet 534, 535, 541 (fig.), 
4 


Simocyoninae, 420 

simplex, Geckobia, 12 
Nicolea, 323 

simulans, Drachylis, 158, 160 

simus, Cyclopidius, 426 

Sinclairella, 434 

singularis, Rymcobites, 617 

Siphonostoma inhabile, 292 
villosum, 290 

Siriella thompsonii, 580 

Sisenes, 88 

sjaelandica, Ctenicera (Malloea), 600 
Elater, 600 

smitti, Ereutho, 328 

snyderi, Amphiprion, 198 

soemmerringii, Gazella, 424 

Solenodontidae, 403 

solitarius, Phorocardius, 608 

sordidus, Vasaces, 87, 89, 92 

Soricidae, 407 

soricina, Glassophaga soricina, 465 

Soricinae, 434 

Soricoidea, 407 

soriniana, Antimelania, 382 

Sorocelides, 171 

Sorocelis americana, 566 

Sparus milli, 198 

speciosissima, Cattleya, 632 

spectabilis, Cirratulus, 286 

see Tapirus, 480, 481, 484, 485, 

486 


Tapirus terrestris, 480, 493 (map) 
Spekia, 371 
zonatus, 371 
spencei, Piezophyllus, 601, 606 
peophila hoffmasteri, 567 (fig.), 569 
(fig.), 570, 571 (fig.) 
spetsbergensis, Eteone, 206, 233, 235 
Sabella, 334 
Sphaeromelania, 366 
largillierti, 365 
Sphaeromias polita, 138 
Sphaeromyas pulchripennis, 137 
Sphaerosyllis, 242, 255 
erinaceus, 207, 243 (fig.), 955 
latipalpis, 256, 257 


661 


longicauda, 256, 257 
Sphaloplana georgiana, 565 (fig.), 566, 
567 (fig.), 569 (fig.) 
Sphenicosomus, 617 
Sphyroperiscelis, 551, 552, 554, 556 
spiloderus, Diploconus, 615 
spinicauda, Chaetura spinicauda, 542 
spiniger, Dennyus, 533, 535 
saa oe Dennyus, 536, 541 (fig.), 
th 


+ 
spinosa, Io, 364 
spinosipenis, Dendrocelopsis, 
Dendrocoelopsis, 177, 
spinosus, Lophogaster, 577 
spinulosa, Lagisca extenuosa, 222 
Polynoé, 217 
Spio, 280, 284 
filicornis, 207, 283 (fig.), 284 
filicornis pacifica, 284 
limicola, 284 
setosa, 284 
Spionidae, 207 (table), 212, 280 (key), 
283 (figs.) 
Spiophanes tenuis, 282 
spirillum, Circeis, 344 
Dexiospira, 344 
Serpula, 344 
Spirorbis, 209, 341 (fig.) 
Spirorbis (Dexiospira), 343, 344 
Spirorbis, 342, 343 (key) 
granulatus, 209, 341 (fig.) 
(Lacospira) granulatus, 343 
quadrangularis, 343 
spirillum 209, 341 (fig.) 


176 
78 


(Dexiospira) spirillum, 343, 344 
spitzbergensis, Sabella, 334 
Eumys, 411, 413, 414 


spokanensis, 
fi 


g. 
Paradjidaumo, 409 
Sponidium, 158 
Squamata, 401 
Stanleya, 388 
neritinoides, 388 
stecki, Beezia, 138 
steenstrupii, Myxicola, 340 
Stelobezzia diversa, 59, 60, 61, 76 
Stenocephaloon, 159 
metallicum, 159 
Stenomelania, 380, 381 
aspirans, 380 
Stenomicra, 551, 557, 560 
angustata, 560 
Stenotrachelus, 159 
Stephanosyllis ornata, 246 
picta, 246 
Sternaspidae, 208 (table), 
313 (figs.) 
Sternaspis, 309 
scutata, 208 
fossor, 209 
islandica, 309 
scutata, 208, 309, 313 (fig.) 
scutata africana, 309 
Sternocampsus, 617 
villosus, 617 


213, 309, 


662 


Stibarus montanus, 430 


sp., 430 
Stilobezzia, 57, 58 (key), 61 (table), 63, 


PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 103 


68, 72 (figs.), 73 (figs.), 135 
(Eukraiohelea) aberrans, 61 
(Eukraiohelea) africana, 63 
antennalis, 59, 60, 61, 68 
(Stilobezzia) antennalis, 68, 73 


(fig.), 83 

beckae, 59, 60, 61 

cu pbenats) beckae, 69, 72 (fig.), 
8 


bicolor, 59, 60, 61, 71 

(Stilobezzia) bicolor, 71, 72 (fig.) 

Biting midges of the heleid genus, 
in North America, 57 

bulla, 59, 61, 74, 83, 84 

(Stilobezzia) bulla, 73 (fig.), 74, 84 

coquilletti, 58, 60, 61, 75 

(Stilobezzia) coquilletti, 72 (fig.), 75 

(Stilobezzia) diversa, 73 (fig.), 76, 


79 
elegantula, 58, 60, 61 
(Eukraiohelea) elegantula, 62, 72 


(fig.) 
(Eukraiohelea) esakiana, 61, 62 
fiebrigi, 76 
fuscula, 58, 59, 61, 64 
(Neostilobezzia) fuscula, 64, 72 


fig.) 
glauca, 59, 60, 61, 77, 85 
(Stilobezzia) glauca, 73 (fig.), 77 
(Stilobezzia) kiefferi, 81 
limnoplila, 83 
lutea, 58, 60, 61, 65, 68 
(Neostilobezzia) lutea, 65, 72 (fig.) 
macfiei, 65 
mallochi, 65, 66 
pallidiventris, 59, 61, 78 
(Stilobezzia) pallidiventris, 78 
paulistensis, 71 
pruinosa, 59, 60, 61, 79 
(Stilobezzia) pruinosa, 73 (figs.), 79 
punctipes, 58, 60, 61 
(Stilobezzia) punctipes, 72 (fig.), 79 
(Stilobezzia) punctulata, 81 
rabelloi, 59, 60, 61, 81 
(Stilobezzia) rabelloi, 72 (fig.), 81 
stonei, 58, 60, 61 
(Neostilobezzia) stonet, 66, 72 (fig.) 
sybleae, 59, 60, 61 
(Stilobezzia) sybleae, 73 (fig.), 82 
thomsenae, 59 
(Stilobezzia) thomsenae, 73 (fig.), 
83 
ugandae, 62 
viridis, 59, 60, 61, 78, 84 
(Stilobezzia) viridis, 73 (fig.) 84 


stone:, Monohelea, 137, 139, 146 (fig.), 


147 (fig.) 

Monohelea (Monohelea), 148, 149, 
150 

Stilobezzia, 58, 60, 61 


Strabala, 121, 122, 123, 124, 125 

(figs.), 126, 127 (key) 

acuminata, 125 (fig.), 127, 128 

acuminata costaricensis, 125 (fig.), 
127, 129 

acuminata teapensis, 125 (fig.), 
127, 129 

ambulans, 125 (fig.), 127, 131 

ambulans jamaicensis, 125 (fig.), 
127,131 

ambulans puertoricensis, 125 (fig.), 
127, 182 

Caen beetles of the genus, 


columbiana, 125 (fig.), 127, 133 
dominicensis, 121 
ferruginea, 122, 125 (fig.), 127, 132 
haematina, 134 
intermedia, 121, 122, 125 (fig.), 
127, 131, 133 
languida, 122 
nigriceps, 122 
rotunda, 125 (fig.), 127, 130, 133 
rufa, 125 (fig.), 126, 127, 128, 129 
rufa floridana, 125 (fig.), 127, 128 
scutellaris, 122, 123 
scutellata, 121 
sp., 134 
trinitatis, 125 (fig.), 127, 134 
Strephobasis, 361 
Streptoprocne zonaris albicincta, 548 
striata, Eteone, 233 
strictus, Neonomopleus, 613 
sire Terebellides, 209, 330, 331 


8.) 
strophensis, Kentrogomphios, 404, 405 
(fig.), 406 (fig.), 430 
Strophitus, 370 
Sturtevant, A. H.; Nearctic flies of the 
family Periscelidae (Diptera) and 
certain Anthomyzidae referred to 
the family, 551 
Stylaroides pluribranchiata, 290, 291 
Subathous, 617 
tonkinensis, 617 
Subathrus, 617 
subglaber, Lophogaster, 577 
sublaevis, Brada, 292 
submetallicus, Hemirhipus, 609 
subnodosa, Nodilittorina, 456 
subterranea, Phagocata, 566 
subtilis, Cosmesus, 601 
Mesembris, 601 
subvittatus, Mullus, 513 
Upeneus, 498, 514, 522, 524 
suifuensis, Melanoides, 380 
suillus, Tapir, 477, 479, 481, 485 
sulcicollis, Pantolamprus, 618 
Sulcilacon, 617 
Sulcimerus, 617 
sulcirostris, Crotophaga, 590 
sulcospira, Melania, 381 
Sulcospira, 381 


Stilobezzia (Neostilobezzia), 66,|Sulcospira, 381 


72 (fig.) 


sulcospira, 381 


INDEX 


sulphureus, Upeneoides, 513, 514 
Upeneus, 500 (table), 501, 502 
(tables), 504 (table), 505 (tables), 
506, 508, 510, 513, 514 (table) 
sulphuripennis, Corymbites, 616 
sundaicus, Upeneoides, 507 
Upeneus, 507, 519 
Supella supellectilium, 40 
supellectilium, Supella, 40 
surinamensis, Blatta, 45 
Pycnoscelus, 45, 51 
sybleae, Stilobezzia, 59, 60, 
Stilobezza Gulabemss). 73 (fig.), 
8&2 


Syllidae, 207 (table), 212, 240, 241 
(key), 243 (figs.), 251 (figs.) 
Syllis, 241, 252 (key) 
alternata, 253, 254 
armillaris, 254 
era 205, 207, 243 (fig.), 252, 
54 
(Ehlersia) cornuta, 253 
fasciata, 205, 207, 243 (fig.) 
(Typosyllis) fasciata, 254 
heterochaeta, 254 
(Ehlersia) heterochaeta, 253 
monilicornis, 260 
oerstedi, 253 
quaternaria, 253, 254 
symmetrica, Oxytrema, 360 
Syndyoceros, 428 
Syspotamus, 477 


Tachornis phoenicobia phoenicobia, 543 
taeniopterus, Upeneus, 507 
Taiwanathous, 617 
arisanus, 617 
Takamatsuia major, 543 
Talpa, 431, 434 
sp., 430 
Talpidae, 403 
Talpinae, 403 
Tanganyicia, 373, 382, 387, 388 
rufofilosa, 387 
tanschaurica, Paludomus, 369 
tapir, Hydrochaerus, 479 
Tapirus, 481, 485, 493 (map) 
Tapirus terrestris, 488, 493 (map) 
Tapir, 477 
(Anta), 489 
americanus, 479 
anta, 477, 479, 481 
maypuri, 479, 481, 485 
pinchaque, 471 
sp., 486 
suillus, 477, 479, 481, 485 
Tapirella, 478, 488 
bairdii, 490 
dowi, 490 
tapirus, Tapirus, 466, 480 
Tapirus, 466, 467, 470, 477, 489 
aenigmaticus, 477, 479, 481, 485 
oe hana. 476, 477, 479, 480, 481, 


americanus mexianae, 480, 481, 485 


663 


andicola, 472 
anta, 481, 485 
antea, 476 
anulipes, 480, 481, 484, 485, 486 
bairdi, 490 
(Elasmognathus) bairdi, 489 
bairdii, 465, 466, 467, 468, 469, 473, 
478, 482 (fig.), 483 (fig.), 488, 489, 
490, 493 (map), 495 (map) 
(Tapirella) bairdii, 489 
bairdii dowii, 491 
brasiliensis, 481, 485 
dowi, 490, 492 
dowii, 480, 484, 490, 491 
enigmaticus, 471, 493 (map) 
ecuadorensis, 479, 481, 485 
excelsus, 468 
indicus, 465, 469 
(Acrocodia) indicus, 467, 489 
laurillardi, 479, 481, 485 
er ane 469, 471, 472, 474, 476, 
4 
Northern Colombia species, 465 
pinchacus, 471, 480 
pinchaque, 465, 466, 467, 468, 469, 
471, 478, 479, 482 (fig. ), 483 
(fig. e 484, 487 (fig.), 489, 493 
(map), 495 (map) 
pinchaque brasiliensis, 479 
roulini, 471, 473, 476, 480 
roulinii, 466, 471, 472, 476 
roulinii brasiliensis, 479 
rufus, 479, 481, 485 
spegazzinii, 480, 481, 484, 485, 486 
tapir, 481, 485, 493 (map) 
tapirus, 466, 480 
terrestris, 465, 466, 467, 468, 469, 
471, 476, 477, 478, 479, 480, 481, 
482° (fig.) 483 (fig. ), 484, 485, 
486, 487 (fig.), 488, 490, 492 
(Hippopotamus) terrestris, 480 
terrestris colombianus, 486, 490, 493 
(map), 495 (map) 
terrestris guianae, P80, 481, 485 
terrestris laurillardi, 479 
eo peruvianus, 479, 481, 
48 
terrestris 480, 493 
(map). 
terrestris tapir, 488, 493 (map) 
terrestris terrestris, 476, 479, 493 
_(map) 
villosus, 471 
Tapyra, 477 
sabatyra, 479, 481, 485 
Tarebia, 378, 379 
granifera, 379, 386 
lateritia, 378, 379 
semigranosa, 379 
tarsalis, Gonodyrus, 610 
tattakensis, Paradima, 614 
Tayussidae, 432 
teapensis, Strabala acuminata, 125 (fig.), 
127, 129 


spegazzinii, 


PROCEEDINGS 


664 


Tectaria montrouzieri, 451 
Tectarius, 449, 450, 454, 455 (fig.), 458, 
461, 462 
coronarius, 461 
muricatus, 449, 450, 452, 453 (fig.), 
454, 455 (fig.), 457 (fig.), 459, 
460, 462 
nodulosus, 462 
tuberculatus, 449, 451, 454 
Tectininus, 458 
telonaria, Melania, 368 
Oxytrema, 368 
temnodon, Paleolagus, 41/5, 430 
Tenalomus, 618 
fulvipennis, 618 
tenebricosa, Hemiblabera, 46 
tenebrosa, Io, 364 
Tenebrionidae, 155 
Tenebrionidea, 155 
tenuicorne, Cephaloon, 157 (map), 158, 
160 


tenuicornis, Corymbites, 614 
tenuis, Euchaetomera, 580 
Maldane, 306 
Petaloproctus, 208, 301 (fig.), 306 
Polycelis, 172 
Prionospio, 282 
Spiophanes, 282 
Terebella maculata, 323 
venustula, 322 
Terebellidae, 209 (table), 212, 318, 319, 
(fig.), 320 (key), 331 (figs.) 
Terebellides, 321, 330 
stroemii, 209, 330, 331 (fig.) 
stroemii japonica, 330 
terebellidis, Saccopsis, 330 
terrestris, Hippopotamus, 479 
Pinchacus, 471 
Tapirus, 465, 466, 467, 468, 469, 
471, 476, 477, 478, 479, 480, 481, 
482 (fig.), 483, (fig.), 484, 485, 
486, 487 (fig.), 488, 490, 492 
Tapirus (Hippopotamus), 480 
Tapirus terrestris, 476, 479, 493 


(map) 
tessellata, Monohelea, 147 (fig.), 150, 
152, 153 
Monohelea (Monohelea), 152 
tessellatus, Corymbites, 600 
Elater, 600 
testacea, Altica, 122 
testaceus, Scarabaeus mobilicornis, 101 
Tetralobus macrocerus, 600, 601 
tecana, Monohelea., 136, 147 (fig.) 
Monohelea (Monohelea), 143 
texana, Geckobia, 15 
Geckobiella, 9, 7/5, 16 (fig.), 17 
(fig.) 
texensis, Chorisoneura, 42 
Thacana, 618 
Thalassema scutata, 309 
Thelepus, 320, 327 
cincinnatus, 209, 327, 331 (fig.) 
circinnatus, 327 
hamatus, 327 
thermarum, Ambrysus, 1, 3 (fig.) 


OF THE NATIONAL MUSEUM 


VOL, 103 


theryi, Jonthadocerus, 611 
Thiara, 378, 379, 386 
amarula, 378 
(Thiara) amarula, 379 
cancellata, 378, 379 
(Setaeara) cancellata. 379 
setosa, 379 
thiarella, Litorina, 451 
Thiaridae, 357, 358, 366, 367, 372, 373, 
374, 875, 377, 378, 886, 461 
Thiaropsis, 378 
winteri, 378 
thompsoni, Domnina, 430 
Eutypomys, 430 
Ictops, 430 
Limnotrochus, 371 
thompsonii, Siriella, 580 
thomsenae, Stilobezzia, 59 
pre Pe (Stilobezzia), 73 (fig.), 
3 


thomsonii, Gazella, 424 
thoracicus, Elater, 605 
tibetica, Polycelis, 172 
tibialis, Nitzschia, 538 
Nitzschia pulicaris, 538 
Tinecus, 618 
Tiphobia, 372, 373, 374 
horei, 373, 374 
Tiphobiidae, 373 
Tiphobiinae, 358, 372, 373 
titanelix, Peratherium, 402, 430 
Titanotheres sp., 430 
Titanotheriomys, 431, 433, 434 
veterior, 408, 430 
tokisensis, ip ees 509 
tonkinensis, Subathous, 617 
torelli, Potamilla, 336 
tornata, Melanoides, 371 
tornatella, Sermyla, 380 
toucheana, Oxytrema, 368 
tragula, Upeneus, 500 (table), 502 
(tables), 508, 504 (table), 505 
(tables), 506, 507, 509, 510, 511, 
514, 521 (table), 622, 524 (table), 
526 (table), 527 
tragulus, Upeneoides, 522 
transmontanus, Leptomeryx, 426, 427 
(table), 430 
Travisia, 295, 296 
brevis, 298 
carnea, 205, 208, 296, 297 (fig.) 
forbesi, 296 
forbesii, 296 
pupa, 296 
tregonboffi, Boreomysis, 579 
Trelasus, 618 
antennalis, 618 
Tremarctos ornatus, 475 
triannulata, Evarnella, 223 
Harmothoé, 222, 223 
Trichobranchus, 321, 329 
glacialis, 209, 329, 331 (fig.) 
glacialis antarcticus, 329 
esa Amphiprion, 189, 182, 196, 
1 


INDEX 


665 


Triclads, Fresh-water (Turbellaria) of | ugandae, Stilobezzia, 62 


Alaska, 163 
tricolor, Amphiprion, 199 
tridens, Boreomysis, 579 
trifasciatus, Amphiprion, 193 
Trigonias, 423 
osborni, 425, 430 
Trigonidiidae, 33 
trilineata, Mudalia, 361 
trilineatus, Autolytus, 250 
Hemirhipus, 605 
trilobata, Ampharete, 316 
trilophus, Paradjidaumo, 409 
trinitatis, Strabala, 125 (fig.), 127, 134 
triocellata, Wetmorella, 441, 442, 443, 
446 (fig.), 447 
tristiculus, Cephaloon variabilis, 161 
Ephamillus variabilis, 161 
Trochidae, 449 
trochiformis, Turbo, 456 
Trochus cumingi, 458 
nodulosus, 451, 456 
Trophonia arctica, 290, 291 
rugosa, 290 
tropica, Maldane sarsi, 304 
truncatum, Pseudomma, 581 
tuberculata, Eteone, 234 
Littorina, 451 
Melania, 375, 377 
Melanoides, 380, 386 


ugiensis, Conobajulus, 606 
Na Tort Oxytrema canaliculatum, 
6 


undulatus, Sceloporus, 9, 15 
unguiculata, Eucopia, 579 
ungulare, Cephaloon, 157 (map), 158, 
159, 160 
unicorne, Bolbelasmus, 97 
unimaculata, Sciaena, 194 
unimaculatus, Amphiprion, 193 
Upeneoides, 506 
arge, 518 
belaque, 513 
caeruleus, 516 
dubius, 514 
fasciolatus, 514 
guttatus, 509 
japonicus, 509 
kiushiuana, 522 
luzonius, 519 
moluccensis, 514 
parvus, 528 
philippinus, 516 
sulphureus, 513, 514 
sundaicus, 507 
tokisensis, 509 
tragulus, 522 
variegatus, 522 
vittatus, 516 


Nodilittorina, 449, 450, 451, 453] Upeneus, 497, 500 (table), 502 (tables), 


(fig.), 454, 455, (fig.), 457 (fig.), 
459, 460, 462 
Pholoé, 230 
Turbo, 451 
tuberculatum, Aylacostoma, 375, 377 
Aylocostoma (Longiverena), 377 
tuberculatus, Litorina, 451 
Tectarius, 449, 451, 454 
Turbo, 458 
tubifex, Eusyllis, 260 
Tubularia sp., 327 
tunicatus, Amphiprion, 191 
Turbellaria, Fresh-water triclads of 
Alaska, 163 
North American triclad, XIII; 
three new cave planarians, 563 
Turbo nodulosus, 451 
Trochiformis, 456 
tuberculata, 451 
tuberculatus, 458 
Turcica, 451 
turgidus, Megalagus, 430 
turriculus, Melanoides, 380, 381 
Tyleudacus, 618 
Tyloarsus, 618 
Tylomelania, 381 
neritiformis, 381 
Tylotarsus, 618 
Typhobia, 371 
typica, Anchialina, 580 
typicus, Lophogaster, 577 
Typitium, 158 
Typosyllis, 261 
alternata, 253 
collaris, 260, 261 


365272—56——4 


504 (table), 505 (tables), 506, 
508 (key), 527 (table) 

arge, 500 (table), 502 (tables), 503, 
504 (table), 505 (tables), 506, 
507, 508, 618 

asymmetricus, 500 (table), 501, 502 
(tables), 503, 504 (table), 505 
(tables), 511, 512 (table) 

bensasi, 500 (table), 502 (tables), 
503, 504 (table,) 505 (tables), 
506, 508, 409, 511 (table) 

bitaeniatus, 516 

bivittatus, 513 

luzonius, 500 (table), 502 (tables), 
503, 504 (table), 505 (tables), 
507, 509, 619, 521 (table), 523, 


526 

moluccensis, 500 (table), 502 
(tables), 504 (table), 505 (tables), 
508, 510, 514 


oligospilus, 500 (table), 502 (tables), 
503, 504 (table), 505 (tables), 509, 
525, 526 (table), 527 (table) 

parvus, 499, 500 (table), 501, 502 
(tables), 503, 504 (table), 505 
(tables), 508, 528 

phillipsi, 498, 528, 529 

Revision of the goatfish genus, 
with descriptions of two new 
species, 497 

subvittatus, 498, 514, 522, 524 

sulphureus, 500 (table), 501, 502 
(tables), 504 (table), 505 (tables), 
506, 508, 510, 513, 514 (table) 

sundaicus, 507, 519 


666 


taeniopterus, 507 
tragula, 500 (table), 
503, 504 (table), 505 (tables), 
506, 507, 509, 510, 511, 514, 521 
(table), 522, 524 (table), 526 
(table), 527 
vittatus, 500 (table), 502 (tables), 
504 (table), 505 (tables), 506, 508, 
510, 516, 517 (table), 518 
(table), 519, 523 
Upenoides moluccensis, 514 
Ursidae, 419 
Ursus horribilis, 419, 424 


vaginata, Dendrocoelopsis, 178 
vaginatus, Dendrocoelopsis, 177 
vainafa, Melanoides, 380 
vanclevei, Caesaromysis, 577, 581 
vandeli, Amyadenium, 177 
Dendrocoelopsis, 178 
vandykei, Cephaloon, 157 (map), 160 
Vannius oculatus, 626 (fig.), 628 
rubrovittatus, 629 
variabile, Cephaloon, 158 
variabilis, Cephaloon, 157 (map), 158, 
159, 161 
Melania, 382 
variegatus, Upeneoides, 522 
Vasaces, 87, 88, 89 (key), 91 (figs.) 
aeneipennis, 87, 88, 89, 91 (fig.), 94 
Beetles of the oedemerid genus, 87 
costatus, 87, 89, 90 
elongatus, 89, 91 (fig.), 93 
knulli, 89, 90, 91 (fig.) 
linearis, 89, 91 (fig.) 
maculatus, 89, 91 (fig.), 92 
sordidus, 87, 89, 92 
vega, Ampharete, 208, 315, 319 (fig.) 
Amphicteis, 315 
Veloplacenta, 383 
maculata, 385 
venusta, Lanassa, 209, 326 
Laphaniella, 326 
venustula, Nicolea, 209, 319 (fig.), 322 
Terebella, 322 
Verena, 376, 377 
verrilli, Autolytus, 246 
versicolor, Eukraiohelea, 61, 62 
Photuris, 35, 36 
verucosa, Pleurocera, 362 
veterior, Titanotheriomys, 408, 430 
vetustus, Prosciurus, 430 
vilis, Litorina, 456 
Nodilittorina, 456 
villosa, Brada, 205, 207, 290 
villosum, Siphonostoma, 290 
villosus, Pinchacus, 471 
Rhino (cheerus), 471 
Sternocampsus, 617 
Tapirus, 471 
violaceus, Pachychilus, 366 
virgatus, Melanoxanthus, 609 
virginianus, Odocoileus, 423 
virginica, Oxytrema, 364 


PROCEEDINGS OF THE NATIONAL MUSEUM 


VOL. 103 


viridipennis, Chaetura chapmani, 545 


502 (tables), | viridis, Ceratolophus, 84 


Ceratopogon, 84 
Hartomyia, 84 
Johannseniella, 84 
Mystides, 232 
Stilobezzia, 59, 60, 61, 78, 84 
Stilobezzia (Stilobezzia), 73 (fig.), 
84 
vitalis, Rostricephalus, 617 
vitalisi, Cardiophorus, 613 
vitida, Raphaea, 616 
vitta, Phagocata, 165, 167 
vittatus, Coracinus, 194 
Elater, 603 
Hypeneus, 516 
Mullus, 506, 516 
Upeneoides, 516 
Upeneus, 500 (table), 502 (tables), 
504 (table), 505 ‘(tables) , 506, 
508, 510 516, 517 (table), 518 
(table), 519, 523 
Viviparidae, 372, 386 
Viviparus, 368, 372 
Vuilletus, 618 


wagenia, Melanoides, 371 

wahlbergi, Paranaitis, 236 

wailesi, Proneomysis, 582 

walli, Pachychilus, 367 

Wanga, 383 

wapleri, Eudactylus, 618 

Wetmorella, 439, 440, (key), 442 (table), 
443 (table), 445, 4.46, 447 

alboraaciat 441, 442, 443, 446, 

44 


ncaligin 440, 442, 443, 444, 446, 

44 

philippina, 439, 440, 446, 447 

philippina bifasciata, 440, 441, 442 
(fig.), 443 

philippina philippina, 440, 441 
(fig.), 442, 448, 445 

Review of the labrid fish genus, 
with descriptions of new forms 
from the tropical Indo-Pacific, 
439 

triocellata, 441, 442, 448, 446 (fig.), 


447 

wheeleri, Periscelis, 552, 553, 556 

White, Theodore E.; Preliminary analy- 
sis of the fossil vertebrates of 
the Canyon Ferry Reservoir 
area, 395 

winnertzi, Microperiscelis, 556 

Periscelis, 556 
winteri, Thiaropsis, 378 
wiréni, Glycinde, 205, 207, 274, 277 


(fig.) 

Wirth, Willis W.; American biting 
midges of "the heleid genus 
Monohelea, 135 

Biting midges of the heleid genus 
Stilobezzia in North America, 
57 


INDEX 


woodburyi, Ambrysus, 3 

worms, Polychaete, from Point Barrow, 
Alaska, with additional records 
from the North Atlantic and 
North Pacific, 203 


Xanthelater, 618 

Xantherater, 618 

Xantholamprus, 618 

Xanthopenthes, 619 

xanthurus, Amphiprion, 190, 192, 198 


Yezodima, 619 
convexuni, 619 


667 


Zalepia, 611, 619 

Zemelanopsis, 357 

ziczac, Littorina, 450, 453 (fig.) 
Littorina (Melarhaphe), 450, 452 

zoea, Gnathophausia, 578 

zonalis, Ellipstoma, 363 
Pleurocera, 363 

zonata, Nereis, 207, 265, 269 (fig.) 
Nereis (Nereis), 264, 265 

zonatus, Spekia, 371 

Zonurobia, 15 

Zorochros, 619 

Zorochrus, 619 

zostericola, Nicolea, 322 


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